ee Laine
teenie: ab eee A
Ratner ere
oak ee Mie en mt Mem kr Sn AS
ed
fon Seb SBOE
<cenianee enil +
viet
Cott ance i a oe 1
To ath eal ete
eeferree
==
Saas
ae as
—
=.
a)
ii
PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
ZAQOLOGICAL SOCIETY
OF LONDON.
1902, vol. I.
(JANUARY—APRIL.)
PRINTED FOR THE SOCIETY,
ANI) SOLD AT THEIR HOUSE IN HANOVER-SQUARE.
LONDON:
MESSRS. LONGMANS, GREEN, AND CoO,,
PATERNOSTER ROW.
Los!
OF THE
COUNCIL AND OFFICERS
ZOOLOGICAL SOCIETY OF LONDON,
1902.
COUNCIL.
(Elected April 29th, 1902.)
His Grace Tue Duke or Beprorp, K.G., President.
Grorce A. Bouencer, Ksq., |
E.R.S. |
Tot Earn or CrAwrorD, K.T.,
ERS:
Wiuiiam BH. pe Winton, Esq.
Hersert Druce, Esq., F.LS.
CuarLtes DrumMonpD, Hsq.,
Treasurer.
Sir Josepu Favrer, Br., F.R.S., |
Vice-President. |
Dr. Cuartes H. Garry, LL.D. |
Dr. Atpert GountHeER, F.R.S., |
Vice-President. |
Carr. THE Marquis or HAmIL- |
ron, M.P. |
Pror. Grorce B. Howes, D.Sc.,
LL.D., F.R.S., Vice-President
Lr.-Cou. L. HowArp Irpy.
Sir Harry Jounston, G.C.M.G.,
K.C.B.
| Sir Huer Low, G.C.M.G.
| P. Cuatwers MircHey, Esq.,
M.A., D.Sc.
| EK. Lorr Puriuips, Esa.
| HowarpSaunpers, Hsq., F.L.S.,
Vice-President.
| Puitie Lutury Sciater, Esq.,
M.A., D.Sc., F.R.S., Secretary.
Dr. Davin Suarp, F.R.S.
OLDFIELD THomas, Esq., F.R.S.
Dr. Henry Woopwarp, LL.D.,
E.R.S., Vice-President.
PRINCIPAL OFFICERS.
P. L. Scuater, Esq., M.A., D.Sc., F.R.S., Secretary.
Frank EK. Bepparp, HEsq., M.A., F.R.S., Vice-Secretary
and Prosector.
Mr. CLARENCE BartLert, Superintendent of the Gardens.
Mr. Arruur Tuomson, Head-Keeper and Assistant Super-
intendent.
Me. F. H. Wateruovuse, Librarian.
Mr. Joun Barrow, Accountant.
Mr. W. H. Coun, Chief Clerk.
Mr. Grorce ArtHuR Dousuepay, Clerk of Publications.
LIST OF CONTENTS.
January 14, 1902.
The Secretary. Report on the Additions to the Society’s
Menasertoum, December WOU - et ansesosn) sscsseenes conc sence
Dr. A. 8S. Woodward, F.R.S. Exhibition of a molar tooth of
aeHossil Elorse Ooi ord tune ste eects. <s sas sane sensat device
Mr. Oldfield Thomas, F.R.S. Exhibition of, and remarks
upon, the skin and skull of a Yellow-backed Duiker
(Cephalophus sylvicultrix) from N.K. Rhodesia............
Mr. W. B. Tegetmeier, F.Z.8. Exhibition of the skin of a
Mountain Hare (Lepus variabilis) which had been
stated to belong to a Hare-Rabbit hybrid..................
1, On Variation in the Number and Arrangement of the
Male Genital Apertures in the Norway Lobster
(Wephrops norvegicus). By F. H. A. Marsuaut, B.A.,,
Christisi Colleges Camillortd se yo. teres seeds acon ecto te ane jane
2. Onsome remarkable Digestive Adaptations in Diprotodont
Marsupials. By Dr. Eivar Lonnsere, C.M.Z.8..........
co
. On the Specimen of the Quagga in the Imperial Museum
of Natural History, Vienna. By Lupwie v. Lorenz,
CNET Ser Coie Bo cantnta sige Nea nels eer ee eee tc ere ot at
4. On a further Collection of Mammals made by Mr. Th. H.
Lyle in Siam. By J. Lewis Bonuorg, M.A. ............
5. On the Insects of the Order Rhynchota collected by
Sir Harry Johnston, K.C.B., in the Uganda Protec-
LOLA HEV AN Verda.) LISTANT ns pees ee Meer hita Ase Gat Sars etl +
Page
bo
12
32
38
iv
6. On two Collections of Lepidoptera made by Sir Harry
Johnston, K.C.B., in the Uganda Protectorate during
the year 1900. By Arruur G. Bururr, Ph.D., F.LS.,
E.Z.8., &c.; Senior Assistant-Keeper, Zoological Depart-
ment, British Museum (Nat. Hist.). (Plate I.) .........
February 4, 1902.
The Secretary. Report on the Additions to the Society’s
Menagerie in January 1902 ..............-.0cet ee ee ete
Mr. F. E. Beddard, F.R.S. Exhibition of, and remarks
upon, the malformed neck-vertebree of a Giraffe that
had died in the Society’s Menagerie
CC eC Ce
Mr. E. Degen. Notice of a Memoir on Ecdysis, as Morpho-
logical Evidence of the original Tetradactyle Feathering
of the Bird’s Fore-limb
1. Notes on the Osteology of the Short-nosed Sperm-W hale.
By W. Buaxtanp Benuam, D.Sc., M.A., F.Z.8., Pro-
fessor of Biology in the University of Otago, New
Healamd: \yi(Plates =) *\ ai ener feces eer cer ace cer aa
2. On a Collection of Dragonflies made by Members of the
Skeat Expedition in the Malay Peninsula in 1899-1900.
By HE. WATpEAW, BA. «(Plates Vir iVale) iaucmaeneece ce
3. List of a small Collection of Orthopterous Insects formed
by Sir Harry Johnston in British Hast Africa and
Uganda in 1899 and 1900, with Descriptions of five new
Species. By W. F. Kirsy, F.LS., FELS., Assistant in
the Zoological Department, British Museum (Natural
History), South Kensington
CC eC eC cc)
February 18, 1902.
Mr. L. W. Byrne, F.Z.8. On the Identity of Lepadogaster
sticlopteryx Holt & Byrne with ZL. microcephalus Brook .
Mr. W. B. Tegetmeier, F.Z.8. Exhibition of, and remarks
upon, the skull of a supposed Hybrid between the Sheep
SNL GIS APUG eos. «av. los vais sce eee Oe RCO ERE REE RECEE
Dr. C. I. Forsyth Major, F.Z.8. Exhibition of, and remarks
upon, some jaws and teeth of Pliocene Voles (JZimomys,
gen. nov.)
Bee mem ee ewer eee ere e er ae reer seserreereseeecereeesesresteone
Mr. Lydekker. Exhibition of, and remarks upon, a skull
and two pairs of antlers of an Elk from Siberia
er reecece
Page
44
51
D4
D4
93
102
Vv
1. On Musiela paleaitico from the Upper Miocene of
Pikermi and Samos. By C. I. Forsyrm Masor, F.Z.S.
(GEIR: WL.) cencinacdisety CoMge eso Rene ee te nann (Um RE
2. On Two new Genera of Rodents from the Highlands of
Bolivia. By Otprrenp Tuomas, F.R.S. (Plates VIII.
5 NID) oon tes Ges acu en ny ae DA rl a
3. On some New Mamuals from Northern N yasaland. By
Oronmenpy MeOMAG HAE Sil esettasnsc cco ce eeckee scoot ee
4, On some Characters distinguishing the Young of various
Species of Polypterus. By G. A. Boutenerr, F.R.S.
Gla GesWe Nir ss ARAMA cis Nene tec rain saute cael. See suas
5. Description of a New Snake of the Genus Psammophis,
from Cape Colony. By G. A. Boutencrr, F.R.S.
CEE PemNCDIN Mana E EE ivi ea em Aa de A, CONES Rtg
6. Observations upon the Carpal Vibrissee in Mammals. By
Frank E. Bepparp, M.A., F.R.S., Vice-Secretary and
Erosector, ofgtie Soctetyy yaaa toc). Suite mores eh ice tide Macaeanes
March 4, 1902.
The Secretary. Report on the Additions to the Society’s
Menagerie in February 1902. (Plate XIII.) ............
Mr. W. B. Tegetmeier, F.Z.S. Exhibition of a series of
photographsiof Prjevalsky’s Elorsel.. fic.......-dece.seeetee
Mx. E. N. Buxton, F.Z.S. Exhibition of a series of photo-
graphic slides illustrative of Bird- and Animal-life on
THlaN@ ys ANY Tan aS) IN FLL oP et Ba a A EOD
Mr. G. T. Bethune-Baker, F.Z.S. Notice of a Memoir on
the Amblypodian Group of the Butterflies of the Family
SORTS OUGKOZ Tor HSPIAR NR nr Gera AND rer Neate ae iy ie
1. On the Origin of Pearls. By H. Lyster Jameson, M.A.,
Bn 7 (Plates XUV =e Vil) icceaaiereaeers es fo. co... s:
2. List of the Parrots represented in the Society’s Collection
in January 1902, with Remarks on some of the Rarer
Species. By P. L. Sctarer, D.Sc., F.R.S., Secretary to
chiey Socienya. (laces) Nav iAlNN i PXSU eee senate acca
3. Descriptions of New Species of Coleoptera of the Family
Halticide from South and Central America. By Marin
JACOB MOSS) Wl CRIES Re, DOC) es Sodas Geniieataremaue ion ade
Page
137
138
138
138
140
166
Ll
vi
March 18, 1902.
Page
Mr, Arthur Thomson. Report on the Insect-house for 1901. 204
Mr. R. E. Holding. Exhibition of, and remarks upon, some
malformed Horns and Antlers ..................02ss0seenees 205
Lt.-Col. J. M. Fawcett. Notice of a Memoir on the Trans-
formations of some South-African Lepidoptera............ 205
1. The Evolution of Horns and Antlers. By Hans Gapow,
WN, IElnD ie IRR Aisi annosodsoscensoccec Gy ae aloha 206
2. On a new Stridulating-Organ in a Scorpion, By R. I.
TR{OCLOGI 1D Adspadonuanbedanencnicacsmocabooconagconare couecoocoouD 222
iso)
. On the Organ of Jacobson in the Elephant-Shrew (Jaero-
scelides proboscideus). By R. Broom, M.D., B.Sc.
(Rib Kae) 0-015) maa nt ooadpoobunenomubooobadbesosanobauianedonsueodacess 224.
4, On some Foraminifera and Ostracoda from Cocos Keeling
Atoll, collected by Dr. C. W. Andrews, 1898. By
FREDERICK CHAPMAN, A.L.S., F.R.MLS..........00.0ceeeeees 228
5, Contributions to the Ichthyology of the Congo.—I. On
some new Fishes from the French Congo. By G. A.
BovuLencer, F.R.S. (Plates XXII.-XXIYV.) ............ 234
April 15, 1902.
The Secretary. Report on the Additions to the Society’s
Menagerie in March 1902. (Plate XXYV.) .............5 237
Prof. F. Jeffrey Bell, F.Z.S. Exhibition of, and remarks
upon, a Starfish with injured limbs which had undergone
VOPALY acc siecewalciajre oae'c aise’ wis aig s Au-otaeenle em talals entatiorcle te erste erase 238
Dr. C. I. Forsyth Major, F.Z.8. Exhibition of, and remarks
upon, some remains of a pigmy Hippopotamus from
GY OTUIS Are ven ns enaialesctecce alae olleiore sole quia pamela tecnico rece eaenerr 238
1. On the Windpipe and the Heart of the Condor. By
Frank E. Beppaxp, M.A., F.R.S., Vice-Secretary and
Prosectororthe Socieby ..., seer eee eee eres 239
2. On the Spiders of the Genus Latrodectus Walckenaer.
By Freperick PickARD CamBRIDGE, F.Z.S8. (Plates
DENG Nag GALL) | Son agn aR er nam indoors soos anndbocunens neon soos 247
vil
Page
3. Notes on the Painted Snipe (Rostratula capensis) and
Pheasant-tailed Jacana (Hydrophasianus chirurgus).
By Frank Finny, B.A., F.Z.8., Deputy-Superintendent
of ‘the Indian Museum, Calcutta. ..5..5...5.00bes.2ccsecesense 261
4, Contributions to the Ichthyology of the Congo.—II. On
a Collection of Fishes from the Lindi River. By G. A.
Bovunencer, F.R.S. (Plates XX VITI.-XXX.) ......... 265
5. Field-Notes upon some of the larger Mammals of Patagonia,
made between September 1900 and June 1901. By
ERE SIGE NE RICE ARID He Zin sc cnniecca ae ticteaias «cama ie ecauese ce 272,
(op)
. Contributions to the Osteology of Birds.— Part V. Yaleoni-
Jormes. By W. P. Pycrarr, F.Z.S., A.L.S. (Plates
PRO NONG IF NO NENSITET (Yi Pe Bi dact ean Meulsminisniaatas ie sletdelericlln aaa 277
AE AY Bek Ti (Ai inal Sih
OF THE
CONTRIBUTORS,
With References to the several Articles contributed by each.
Brepparp, Frank E., M.A., F.R.S., Vice-Secretary and
Prosector to the Society.
Exhibition of, and remarks upon, the malformed neck-
vertebre of a Giraffe that had died in the Society’s
IVC MAG OTC awl gsr (elon eran = qaemseitas a niga wh aoa iva pe sMiayiny, at
Observations upon the Carpal Vibrissee in Mammals ...
On the Windpipe and the Heart of the Condor .........
Bett, Professor F, Jerrrey, M.A., F.Z.8.
Exhibition of, and remarks upon, a Starfish with
injured limbs which had undergone repair ...............066
Benuam, W. Buaxiann, D.Sc., M.A., F.Z.S., Professor of
Biology in the University of Otago, New Zealand.
Notes on the Osteology of the Short-nosed Sperm-
Wihallets se (Blates Bl TV: ew was uud a staisns sis cenes sme sl
Page
238
a4
Beruvune-Baxer, G. T., F.Z.8.
Notice of a Memoir on the Amblypodian Group of the
Butterflies of the Family Lycenida
Bee eet ore eee e reese nereoesene
Bonnots, J. Lewis, M.A., F.Z.S.
On a further Collection of Mammals made by Mr. Th.
H. Lyle in Siam
foe eee eee eee oes ec reeses so soseeeeeesessteesneenseaese
BouLEeNcerR, GEorcE AtpBert, F.R.S., F.Z.8S.
On some Characters distinguishing the Young of various
Species of Polypterus. (Plates X. & XI.) ...............40
Description of a New Snake of the Genus Psanmophis,
EromiCape Colony (Plate SXells\ renee. case heen eee neee eee
Contributions to the Ichthyology of the Congo.-—I. On
some new Fishes from the French Congo. (Plates
XXIT.-XXIYV.)
Pee meee ere reeseeenesrersceeoeseereesereessseseresen
Contributions to the Ichthyology of the Congo.—II.
On a Collection of Fishes from the Lindi River. (Plates
XXVITI.-XXX.)
Peewee e erase rete sete r essere sesesesseesesesessesee
Broom, R., M.D., B.Sc., C.M.Z.S., Pearstown, 8. Africa.
On the Organ of Jacobson in the Elephant-Shrew
(Macroscelides proboscideus). (Plate XX1.)
wee sere se eeeeee
Buruer, ArtHur G., Ph.D., F.LS., F.Z.8.
On two Collections of Lepidoptera made by Sir Harry
Johnston, K.C.B., in the Uganda Protectorate during
Wave een ISTO (MEd n We RIES Pema deeosoconocc:obsnobococosesanonnance
Buxton, Epwarp Norts, F.Z.S.
Exhibition of a series of photographic slides illus-
trative of Bird- and Animal-Life on the White Nile ......
138
38
234
265
224
44
Xl
Page
Byrneg, L. W., F.Z.8.
On the Identity of Lepadogaster stictopteryx Holt &
Byrne with Z. microcephalus Brook .............cccsceeeeeenes 102
CAMBRIDGE, FREDERICK PICKARD, F.Z.S.
On the Spiders of the Genus Latrodectus Walckenaer.
(GBllates RONG TE EXENG VAT nicer solhnu ai savers iceuagndes yee 247
CHAPMAN, FrepDERIcK, A.L.S., F.R.M.S.
On some Foraminifera and Ostracoda from Cocos
Keeling Atoll, collected by Dr. C. W. Andrews, 1898 ... 228
DecEN, Epwarpb, F.Z.S8.
Notice of a Memoir on Eedysis, as Morphological Evi-
dence of the original Tetradactyle Feathering of the
TESHEG HS) LNGIRS hie Chae rM eaten Who C Me mR eR en mmr Nate Cima METAS hE ar 54.
Distant, W. L., F.E.S.
On the Insects of the Order Rhynchota collected by
Sir Harry Johnston, K.C.B.,in the Uganda Protectorate. 41
Fawcert, Lt.-Col. J. M., 5th Lancers.
Notice of a Memoir on the Transformations of some
Sout Acme came pid OMLCLAN I «cais ass caes ar iate cca seas siecle 205
Finn, Fran, B.A., Deputy-Superintendent of the Indian
Museum, Calcutta.
Notes on the Painted Snipe (Lostratula capensis) and
Pheasant-tailed Jagana (Hydrophasianus chirurgus) ...... 261
GaAbow, Hans, M.A., Ph.D., F.R.S., F.Z.8.
The Evolution of Horns and Antlers .................. an POG
Houpine, R. E.
Exhibition of, and remarks upon, some malformed
loves: yet! Je Wai) Kerk Ragangneas anc nd ane mano acboddueessicobaconcce ts 205
Xli
Jacosy, Martin, F.E.S.
Descriptions of New Species of Coleoptera of the Family
Halticide from South and Central America. (Plate XX.)
Jameson, H. Lysrur, M.A., Ph.D., Municipal Technical
College, Derby.
On the Origin of Pearls. (Plates XIV.-XVIL.) ......
Kirsy, W. F., ¥.LS., F.E.S., Assistant in the Zoological
Department, British Museum (Natural History),
South Kensington.
List of a small Collection of Orthopterous Insects
formed by Sir Harry Johnston in British Kast Africa
and Uganda in 1899 and 1900, with Descriptions of five
MLO WIS PCCLOS ai sciciciematicielvinsericionusmian abe tem actee alee aceeee nee Reet
Larpiaw, F. F., B.A., Assistant Lecturer and Demonstrator
at Owens College, Manchester.
On a Collection of Dragonflies made by Members of the
Skeat Expedition in the Malay Peninsula in 1899-1900.
(lattes AV ids Vals) vie ie ode ode AUS nine dentaeraciome tenes
Lonnpere, Dr. Ernar, C.M.Z.S8., of the University, Upsala.
On some remarkable Digestive Adaptations in Dipro- _
focomt: Miarsipials’ isc cclois ewes ae incre we wae en a tlcle aa laa Ree nteee
Lorenz, Lupwic von, C.M.Z.8., Imperial Museum of
Natural History, Vienna.
On the Specimen of the Quagga in the Imperial Museum
of Natural History, Vienna ‘a: J2.esteeesseescaeer seen en ecees
LypEekkeEr, R., B.A., F.R.S., F.Z.S.
Exhibition of, and remarks upon, a skull and two pairs
Otantlers of an (Blk trom: Silberiasse-cceeeespeeeaeeeee eee
Page
171
140
93
(Su)
bo
xiii
Masor, Dr. C. J. Forsytu, F.Z.8.
Exhibition of, and remarks upon, some jaws and teeth
of Pliocene Voles (MWimomys, gen, NOV.) ............cceeeeee
On Mustela paleattica from the Upper Miocene of
Pikermuand samosy “(Plate VIE isis sdecdsess aces tes ges
Exhibition of, and remarks upon, some remains of a
igmy Hippopotamus from Cyprus ..................ieecees
[OUST ASS OTD:
MarsHat., F. H. A., B.A., Christ’s College, Cambridge.
On Variation in the Number and Arrangement of the
Male Genital Apertures in the Norway Lobster (Wephrops
MORVEGUCUS sean essed PIERS Oe ee hd a se Ata ah es ae oa en
Pocock, R. I., F.Z.8.
On a new Stridulating-Organ in a Scorpion ............
PricHarD, Heskety, F.Z.S.
Field-Notes upon some of the larger Mammals of Pata-
gonia, made between September 1900 and June 1901
Pycrart, W. P., F.Z.S., A.LS.
Contributions to the Osteology of Birds.—Part V.
Falconiformes. (Plates XXXI.-XXXITIL) ...............
Scuater, Puiu Lurury, M.A., D.Sc. Ph.D., F.RBS.,
Secretary to the Society.
Report on the Additions to the Society’s Menagerie in
ecommerce OOM oe assoc cyan ab er en a eta ae yer ene
Report on the Additions to the Society’s Menagerie in
January 1902
Report on the Additions to the Society’s Menagerie in
February 1902. (Plate XIII.)
List of the Parrots represented in the Society's Col-
lection in January 1902, with Remarks on some of the
Rarer Species. (Plates XVIII. & XIX.)
Report on the Additions to the Society’s Menagerie in
March 1902, (Plate XXV.)
Pam e em eer eee reese see eeneseeeesssesssrsereeserssverece
CR Oe Oe i
Cee meres eee sree reer eersereessone
109
238
bo
277
xiv
TEGETMEIER, W. B., F.Z.S.
Exhibition of the skin of a Mountain Hare (Lepus
variabilis) which had been stated to belong to a Hare-
Vez) eV Hal 0716) GIN t Nata amir Sry SRA SOR Ree RANE NPRPER AAD SAE cU.5 S00
Exhibition of, and remarks upon, the skull of a supposed
Hybrid between the Sheep and the Pig .....................
Exhibition of a series of photographs of Prjevalsky’s
fs (OT EST Stee RAI Near ney aA AE Me gs BSR Gs Tal aoe ORNS 8
THomas, OLDFIELD, F.R.S., F.Z.S.
Exhibition of, and remarks upon, the skin and skull of
a Yellow-backed Duiker (Cephalophus sylvicultria) from
DINAH) EVN OG CSTE: 3. cso t okt" catsieietc os arelct oto erelaes cane aeRO
On Two new Genera of Rodents from the Highlands
Of Bolivia: (Plates ev aiMlindslXe) |e eeeperas see ee ere ae eee r
On some New Mammals from Northern Nyasaland
Tuomson, ArtHuR, Assistant-Superintendent and Head-
Keeper of the Society’s Menagerie.
Report on the Insect-house for 1901 ..................005
Woopwarpb, ArtHur Smuiru, LL.D., F.R.S., F.Z.8.
Exhibition of a molar tooth of a Fossil Horse, Ono-
LOU UE ss doo hea oui SRO oad ohne qe Pve cian eee an en
Page
204
LIST OF PLATES.
1902.—Vot. I.
Plate
lepeWepidopterayirnommUeand ay syria alesse soleil:
UTE)
IUDL, | Osweallnegy Gi Cage -ocoadadcnagbooonpsnascsecne
IV.
oe Dragonflies from the Malay Peninsula............
VII. Skulls and Teeth of Mustela paleattica ..........
VIII. Neoctodon simonst
oe eee woe eee eae re toes eee eee ae e
IX. 1,2,3,44,& 6a. Andinomys edav, adult; 46& 66.
Young of do.; 5a& 7a. Chinchillula sahame,
adult; 56 & 76. Young of do. ; 8-12. Neoctodon
SEPOUIST AAU bina apt costtet arel Mevor aie sinetaesens soci) oie
X. 1,2. Polypterus lapradi. 3. P. weeksti ........
XI. 1. Polypterus congicus. 2. P.endlichert. 3. P. ve!
GARE, 8s IR OTOES oop op oo05 HOOF oe Bl p55 20
SOUS JASONS UBGTURUD ob ocadinnos cocoon adcusooe
OUD, LIS FAI 5 Shoo ocnooo dec duo agecsu ou mano:
XIV.
ai Jameson on the Origin of Pearls ........2.....4.
XVII.
SNAUOL, DORIGCHIS CRCRUGRTOGHON, 6p Boaseccobooosouuoueoo
SKIEXG © Plat Cen cus) MUASTENSTANUS) oh Vance 1. oon ns «os
OSG INGGT STSGIOS Cl LEIGH ooocngeonoanseoongudone
XXI. Jacobson’s Organ in Maceroscelides
CC et
XXII. 1. <Adlabenchelys longicauda, 2. Clariallabes melas. .
ONT Graben lelculeie a ie perth donde ci eiecis cc sieeve
OMY, CRLOAROTS CITED, os 00 aceobgnboonoe EDD OUUODD
XXV. Cercopithecus otoleucus ..cc..cev veers see tnenes
oe Spiders of the Genus Latrodectus....,...+.05++..
Page
Xv1
Plate Page
XXVIII. 1, 2. Micralestes stormst. 3. Phractura lindica ..)
XXIX. 1. Auchenoglanis punctatus. 2. Auchenoglanis a
GUGPs Sin, BUDO GITES OREOES ao nbe bese scouocce > 265
XXX. 1. Pseudoplesiops squamiceps. 2. Tilapia storms. |
Bibs d RUSICHTI PR CHUA We 5 ogo pace isa ao4 oo J
XXXII.
XXXII.> Osteology of the Fulconiformes ....... 0.00000 cee 277
XXXII.
LIST OF TEXT-FIGURES.
1902.—Vot. I.
Page
1. Genital apertures of Nephrops norvegicus ...... iyo Hai ON 4
2. Genital apertures of Nephrops norvegicus ....0.eeseeve creas 4
3. Diagram illustrating variation in genital apertures in Nephrops
MOTVEGECUS oo. ecenenseseeres Nickens a hserarcvare specone everate
4, Ceecum of Trichosurus velpecula oo... cv ve cece eee ee een nees 16
5. Ceecum of Pseudochirus ccetdentalis .......ss sees Pores tes)
6. Ceecum of Peéawrus breviceps 1... .c cere cne cere ene ee snes 24
7. The Quagga of the Vienna Museum .............s-+05- Sieve oe
8, Odontopus notabilis ...seeeecsv eee ces ees ees cen tin they orc 43
9, Cervical vertebrae of a Giraffe ...... ERIS OSE Silage Car euctcar' 53
10. Third femur of Tetrathemis hyalinia ....... Sees, ep aaah e aes a
11. Second femur of Zygonidia malayand ......ve1005. Sienciene ieee 74
12. Third leg of Onychothemis testacea ...... IWndine dt > uo OE On 76
13. Teeth and jaws of Tertiary Voles ....,..-0+.sseeeeeeeeeee 108
14, Teeth of Tertiary Voles, enlarged ..... 2.0. eee e eee eee eee 105
15. Teeth of Voles from Forest Bed and Norwich Crag ........ 106
16. Skull and antlers, with the upper cheek-dentition, of Siberian
IDS A oige Oo cep IEE DUO n seve ttonircusvers ® sor siento 108
17. Left fore foot of Dasypus villosus, ventral surface .......... 128
18. Right fore foot of Petaurus sciwreus, ventral surface ....... . 180
19. Right hind foot of Petawrus sctwreus, lateral surface ...... So ee
20. Left fore foot of Nasua narica, ventral surface .............. 1382
21. Left fore foot of Hyrax, ventral surface .........5- Deretannrae 134
22. A Pearl about to become attached to the Shell ............ 150
93. Cuticle of the Cercarta, in surface VIEW... ww eee ee eee vO
DA. Tapes decussatus .viveseecvecereverenes ori icrialescvtete Snot
25. Evolution of Horns and Antlers .............. iqodsor. woaico ses G
26. Stridulating-organ of Parabuthus flavidus ........ Pron OR 223
27. Cytheridets andrewst ....ccc cece cece nner tn ene e enn eees 229
28. Cytherella vesiculosd ....ccr cern eeeree ee ee ce ee ence nennnes 23
29. Lower end of windpipe of male Sarcorhamphus gryphus, front
WON. . oesuduoududBahouss oo Gh oogGdodgoodn deco Obcod 240
Proc. Zoou. Soc.—1902, Vou. I. b
36.
37.
XV1il
. Lower end of windpipe of male Sarcorhamphus gryphus, back
BVT WW) Ss eure ns favre ciate acre Ge nt tntel he ld er ce
. Heart of Sareorhamphus gryphus opened so as to display the
INtenioMmolthe meh ventriclewe.., ni. ier eee
. Heart of Scythrops nove-hollandie cut open so as to display
interornior melt wentricle sn. nays cet cer eee
. Left lateral aspect of the sternum and havi laneariills of Ser-
pentarius serpentarius, showing the articulation of the
furcula withathe caring graenienic cece serene
. Left lateral aspect of the sternum and shoulder-girdle of Aquila
TAPAL wvcceves eoeceteeaee e@oeeeeoevee o ooeeveeoe? ® eee eee @eee
. Dorsal aspect of the sain of Prantagemaes californianus,
showing the Ciconiine character of the pelvic girdle ......
Dorsal aspect of the pelvis of Cathartes aura ........ acne
Dorsal aspect of the pelvis of Pandion haliaétus ..........0:
Page
241
244
245
297
299
LIST OF NEW GENERIC TERMS
PROPOSED IN THE PRESENT VOLUME (1902, vot. I).
Page
Allabenchelys (Pisc.) ............... 234
Andinomys (Mamm.) ............ 116
Chilochromis (Pise.) ............... 236
Climacobasis (Neuropt.) ......... 85
Page
Oryptomima (Lepidopt.) ........- 50
Mimomys (Mamm.) ............++ 102
Nasigona (Coleopt.) ........s0..00. 203
Neoctodon (Mamm.):....,.... vee Le,
a ys /
sees
“head
Be AMP EWG)
: acai
be ba.
OHI AID, WMA Te) PeLde
wilt LOFT abana Ue Lee
rN :
(aleobigss MPR N YO RRR io) ae SAE
Cintue lt) ayposuilt EP ikea. RD SOR eee
egn eeeysaa es oh as Kb "1h lear he nA ; B89 Pune keis aes is (aud ngaveernalnadin’ J
BA dee) Dario el 3) canned ces hi degh bi) sido Sy
Li
PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.
1902, Vol. I. (January to April).
January 14, 1902.
Prof,.G. B. Howss, LL.D., F.R.S., Vice-President,
in the Chair.
The Secretary read the following report on the additions to the
Society’s Menagerie during the month of December 1901 :—
The registered additions to the Society’s Menagerie during the
month of December 1901 were 51 in number. Of these 11 were
acquired by presentation and 4 by purchase, 6 were born in the
Gardens, and 30 were received on deposit. The total number
of departures during the same period, by death and removals,
was 146.
Dr. A. S. Woodward, F.R.S., exhibited a newly-discovered
upper molar tooth of a Fossil Horse, Onohippidium, from the
cavern near Consuelo, in Last Hope Inlet, Patagonia’. This
specimen was fixed in the bone and carried traces of the soft
parts.
Mr. Oldfield Thomas, F.R.8., exhibited the skin and skull of a
female Yellow-backed Duiker (Cephalophus sylwicultrix) which
orale 74 Sp VEO 0s 1
Proc. Zoou, Soc.—1902, Vou. I, No. I. 1
2 MR. F, H, A, MARSHALL ON [Jan. 14,
had been obtained by Mr. B. B. Johnstone (Native Commissioner)
in the Awemba district of North-eastern Rhodesia, and had been
sent home and presented to the British Museum by Mr. Robert
Codrington, Administrator of that country.
This specimen proved a very considerable extension of the range
of the species, hitherto known only from West Africa, where it had
a wide distribution, from Sierra Leone to Angola. No differences
of importance, however, were perceptible between the Rhodesian
specimen and examples from West Africa.
Since Mr. Codrington’s specimen had arrived, a frontlet and
piece of skin of the same species had also been received from
Mr. O. Baragwanath of Bulawayo, and this, it was believed, had
been obtained in the same district.
In N.E. Rhodesia this Antelope was said by Mr. B. B. Johnstone
(who had obtained the specimen for Mr. Codrington) to be met
with in stony localities at fairly high altitudes. It was sup-
posed to occur throughout the Luemba Highlands and along the
Mohinga Mountains east of Lake Bangweolo, but was not
common. Its cry was like that of a Duiker. Its native name
was “Chibusimawe ” (=Big Mountain Goat).
Mr. W. B. Tegetmeier, F.Z.8., exhibited the skin of an animal
which it had been suggested was a hybrid between a Hare and
a Rabbit, but which had proved to be merely a Mountain Hare
(Lepus variabilis).
Mr. Tegetmeier also exhibited a skull of a Rabbit showing
overgrown incisors in both jaws.
The following papers were read :—
1. On Variation in the Number and Arrangement of the
Male Genital Apertures in the Norway Lobster
(Nephrops norvegicus). By F. H, A. Marsnatt, B.A.,
Christ’s College, Cambridge *.
[Received November 21, 1901.]
(Text-figures 1-3.)
The total number of specimens of the Norway Lobster examined
for the purposes of this investigation was 1123, of which 1080
were males. The latter presented no less than ten different
arrangements of the genital apertures, in addition to the normal
arrangement of an opening on the basal joint of each of the fifth
walking-legs, Before giving the numerical details of the degrees
1 Communicated by W. BarEson, F.Z.S.
1902. | VARIATION IN THE NORWAY LOBSTER. 3
of frequency of these arrangements, it will be convenient te
describe each of them separately. They are as follows :—
A. The normal arrangement.
B. The normal apertures and two additional ones at the
bases of the fourth legs, making a total of four
apertures.
C. The normal apertures and an additional one at the base
of the left fourth leg, making a total of three apertures.
D. The normal apertures and an additional one at the base
of the right fourth leg, making a total of three
apertures.
EK. The normal apertures and four additional ones on the
third and fourth legs, making a total of six apertures.
F, The normal apertures and three additional ones on the
fourth legs and the left third leg, making a total of
five apertures.
G. The normal apertures and three additional ones on the
fourth legs and the right third leg, making a total of
five apertures.
H. The normal apertures and two additional ones on the
left third and left fourth legs, making a total of four
apertures.
K. The normal apertures and an additional one on the left
third leg (there being a gap in the series), the total
number of apertures being three.
L. The normal apertures and two additional ones on the
right third and left fourth legs, the total number of
apertures being four.
M. The normal apertures and five additional ones on the
fourth and third legs and also on the left second leg,
the total number of : apertures being seven.
These arrangements may be represented graphically as follows :—
B. =~ Op D. EK.
r. lL. r. l. Te d. Tr l.
Third legs. . :
° : - Fourth legs. - . °
° : : - Fifth legs. : ° : °
F G lel, K
r l. r l Tes l 7 L
° . Third legs, ° .
. . : - Fourth legs. °
° ° ° - Fifth legs. 2 : ° °
L M.
r L. r L.
Second legs, °
° Third legs. . .
« Fourth legs. - °
. - Fifth legs. . :
[The letters 7. and /, denote the right and left sides. |
he
MR. F. H. A. MARSHALL ON
Text-fig. 1.
[Jan. 14,
Genital apertures of Nephrops norvegicus.
Male of Nephrops norvegicus having abnormal genital apertures on each of the
third and fourth walking-legs.
Text-fig. 2.
Genital apertures of Nephrops norvegicus.
Male of Nephrops norvegicus having abnormal genital apertures on the right third
and right fourth walking-legs.
In no specimen was either of the normal apertures wanting.
The animals were examined in batches—the first batch, which
was much the largest, consisting of the stock of Norway lobsters
in the zoological laboratory of the University of Edinburgh,
obtained for the use of students during the summer session.
The rest were procured at various intervals of time during
the summer and autumn,
are now given :—
(1) Females
Pee S eres es eerererscereseesesesessessece
Normal males
Came score eeee reser eeseseeee sesso
Abnormal males with arrangement B...
Ditto with arrangement C
29
thd
9
eee eer ee res eeee
feo eer essere sre
The numerical details of these batches
1902.]
(2)
(3)
(4)
(5)
(6)
VARIATION IN THE NORWAY LOBSTER.
Females ...............
Normal males
Pee eC a eC Ce Cay
Abnormal males with arrangement B...
Ditto with arrangement C
9
”
Females ...............
Normal males.........
weet ee ewe eres
wee e eee e eee s eee
Peewee eee e renee ere cecene
ee ay
Abnormal males with arrangement B...
Ditto with arrangement C
”
”
IBlemailesieern eee sen:
Normal males
Ce ee
eee ese eee e eee
Total ...
Abnormal males with arrangement B...
Ditto with arrangement C
Females ...............
Normal males
ecco reese eceeee
cere reer eee cere
eee reece ees oeee
eC ee
wee e eter reer ere e rete sess
Abnormal males with arrangement B...
Ditto with arrangement C
Females ...............
Normal males
wee e creer eee eee e seers eeee
seem terre eee teense ecene
Abnormal males with arrangement B...
Ditto with arrangement D
D
60
3) |
1 67
apt r
Voy
69
3
80
5)
3
SGU
2 |
ih as
97
4
95 )
1)
Syl aa
5 gig) ae
1
ie
112
1
Ang 8
2)
1
2 | A
Dane :
1
re
50
1
29
1 24
ia]
6 MR. F. H. A. MARSHALL ON [Jan. 14,
(ia eubiemm all ostyc on. eter testi BOS UL, Raed Sch 1
iINormealltimailes:. ass savesteae maces sads asset (6)
Abnormal males with arrangement B... 4 85
Ditto with arrangement C ............... 5 > 12
“5 * UD) Se osenaee ns eme 3
Total . 86
(8) se Nornaaillenall esteeraeieee se ance c cane 22
Abnormal males with arrangement B... 1 25
Ditto with arrangement C ............... ts &
as 63 LD Pree aen ee 1
Total 25
Adding all these together we arrive at the following result :—
Memeo sale Mod. ated wan. vee hanes a eee 68
Normal sialles art, enen scene aay 878
Abnormal males with arrangement B... 40 )
Ditto with arrangement C ............... 40 |
ri 3 dD See Re Aan 31 |
js 5. A eer orl trcye 2
: ve 1 Flee a ae 3 1000
i GP tapeap a Ve ae
5 *, EL oe ee 1
‘ % LESGRED -nciens a 1
5 i Da PE ee diesnantee if
zs IN Desi. See Loe 3)
Grand Total... 1068
The total number of male Norway lobsters examined being
1000*, the percentage of abnormality occurring among them is
shown to be 12:2. The numerical variation in the apertures, but
not the variation in their arrangement, I have indicated by a
percentage curve (text-fig. 3, p.7). It is of interest to note that
the homeeosis occurs with little regard to bilateral symmetry.
The relative scarcity of females is worthy of comment. It may
be that the majority of them had migrated toa greater distance
from land. The 68 specimens that were examined possessed only
the normal apertures on the third pair of walking - legs.
Dr. Malcolm Laurie, however, tells me of a female specimen in
his possession which has two pairs, the additional ones being on
the fourth pair of legs.
The 1068 Norway lobsters which enter into my calculations
1 Since the above was written I have received 24 male Norway lobsters from the
Forth area, 21 being normal, one showing arrangement C and another arrangement
D, while a third presented an arrangement not hitherto observed, having apertures
upon the left third and right fourth walking-legs in addition to the normal ones,
the total number of apertures being four.
1902. | VARIATION IN THE NORWAY LOBSTER. 7
were all obtained from the area of the Firth of Forth. In addi-
tion to these I procured 80 specimens which were caught off the
Isle of Man. Of these two were females, one was an abnormal
male having two pairs of genital apertures, and the rest were
_ Text-fig 3.
Diagram illustrating variation in genital apertures in Nephrops norvegicus.
Percentage-curve illustrating variation in the number of genital apertures in 1000
male specimens of Nephrops norvegicus. 'The lower figures give the number of
apertures and the side figures the percentage of individuals.
normal males. The number is, of course, not large enough for
any definite conclusions to be based upon it; but in view of the
fact that I never obtained even smaller batches from the Forth
area without finding a much higher percentage of abnormality,
the presence of only a single abnormal specimen among the Isle
8 MR. F. H. A. MARSHALL ON [Jan. 14,
of Man lobsters may point to the percentage being related to the
locality *.
The positions of the additional openings upon their respective
legs are approximately the same as those of the normal ones on
the fifth legs. In the case of those specimens with three pairs of
openings, the most anterior of which are situated on the third
legs in the position of oviducal openings, it is clear, if only from
the modification of the anterior abdominal appendages, that the
Specimens are males. The abnormal apertures are sometimes
smaller than the normal ones, though they may be even larger.
In the case of the single specimen showing seven spermatic
apertures, the six posterior openings are of about equal size,
while the opening on the second thoracic leg on the left side is
very much smaller but still quite obvious.
The state of preservation of the majority of the specimens
rendered it impossible to determine the structure of the internal
genital organs. In the fresh specimens it could, however, in some
cases be made out that the apertures opened internally into blind
sacs. In a few there appeared to be duct-like extensions of these
sacs internally. In a fair proportion the vasa deferentia gave
off branches which extended for a short distance towards the
abnormal openings. In at least one instance these anterior forks
of the vasa deferentia reached the bases of the legs on which the
abnormal apertures were situated. Whether there is ever a free
functional passage from the position of the forking to the
abnormal aperture it is difficult to say with certainty.
That Norway lobsters with additional genital apertures have
been common in Scottish waters for a considerable number of
years, appears from information supplied me by Professor Ewart,
Dr. Beard, Dr. Masterman, and others. Before I began my
investigation on the degree of frequency of such abnormal
lobsters, Dr. Masterman expressed the opinion that quite 10 per
cent. of the specimens he had observed since he had been in
Scotland had additional genital openings; and Dr. Beard, who
has had occasion to examine a very large number, speaks to me
of regular epidemics of this kind of abnormality in some years
in the past, the students in the laboratory experiencing great
difficulty in distinguishing the males from the females.
The only published record, so far as I know, of additional
genital openings in WVephrops is a recent paper by Mr. Cole, who
states that “abnormalities in oviducal and spermatic apertures
are by no means uncommon, and I remember examining three
specimens, two of which were abnormal and had four super-
numerary spermatic apertures occurring as follows :—
Third walking-legs. «
: - Fourth walking-legs. -
. ¢ Fifth walking-legs. : :
1 Mr. Bateson informs me that he has noticed some variation in the degree of
frequency of abnormality in regard to the oviducal apertures in batches of Astacus
procured at various times for the Zoological Laboratory at Cambridge, and is of
opinion that this variation is probably related to the localities from which the
batches were obtained.
”?
1902. ] VARIATION IN THE NORWAY LOBSTER. 9
Mr. Bateson has, however, placed on record several cases of
females of Astacus fluviatilis with additional oviducal apertures,
but their degree of frequency was not nearly so great as that of
the abnor anal spermatic apertures in Wephrops. After citing
Desmarest’s observation of a female Astacus with oviducal
apertures on both the antepenultimate and penultimate legs, to
each of which the oviducts branched, he describes several cases
that he has himself observed. Among 583 female Astaci he
records 23 which were abnormal in regard to the genital aper-
tures, 17 having an opening on one of the fourth legs, one with
an opening on cach of the fourth legs, one with one opening on
each of the fourth and fifth legs (in “each case in addition to the
normal openings), and four in which one of the normal openings
was wanting. The oviducts in most cases gave off branches to
the abnormal openings as in Desmarest’s specimen. Mr. Bateson
cites Dr. Benham’s observation on a female crayfish which had a
pair of supernumerary openings on the fifth legs but none on the
fourth. Out of 714 males that Mr. Bateson examined, one was
abnormal in having no spermatic aperture on the right side. No
cases of additional spermatic apertures are recorded for Astacus.
The above-described variations in Nephrops would appear to
have some bearing on the supposed cases of hermaphroditism
among the Astacide. La Valette St. George has described a
specimen of Astacus fluviatilis, in its external characters a male,
but with what appeared to be a hermaphrodite gland. Bergendal
in two papers has recorded his observations on females of Astacus
fluviatilis in which the appendages of the first abdominal somite
were modified as in the male; and Faxon has cited other cases of
partial or complete hermaphroditism. But it is only those cases
where the evidence of hermaphroditism is supplied by the exist-
ence of apertures situated as in one sex, inanimals which in many
characters resemble the other sex, which specially concern the
subject of this paper.
Tn his ‘ Revision of the Astacide’ Faxon gives an account of a
specimen of Cambarus propinquus, which appears to have been an
undoubted female, for ovarian eggs were found on dissecting it.
The external characters, including the condition of the appendages
of the first and second abdominal somites, were also those of the
female, with the exception of the position of the genital apertures,
which were on the last pair of thoracic legs—z. e., in the position
typical of the male.
Lénnberg states that he believes he has seen rudimentary
genital ducts passing to the third pair of thoracic legs in two
specimens of Cambarus fallax, but owing to their state of preser-
vation he is not positive.
Von Martens has long ago recorded the presence of additional
apertures on the bases of the antepenultimate legs in certain
male specimens of Cheraps preissii, Astacus pilimanus, and
A. brasiliensis, the two latter of which are now included in
Huxley’s genus Parastacus.
‘Von Ihering describes these apertures which occur in all the
10 MR. F. H. A. MARSHALL ON [Jan. 14,
specimens of Parastacus he saw as follows:—‘“ Il y a sur le coxo-
podite de la troisiéme jambe, une ouverture ovale qui est fermée
par un écusson bombé et que l’on peut déprimer du cdté médian
ou libre.” The state of preservation was not good, but von Ihéring
says :—‘‘Tl m’a paru qu’un conduit trés délicat se dirigeait plus en
avant, & louverture du troisiéme coxopodite, mais je ne puis
Vaffirmer.” Whether or not the specimens dissected were her-
maphrodite, von [héring is apparently also doubtful.
Faxon records the coexistence of both pairs of apertures in all
the specimens he has examined of Parastacus saffordi, P. vari-
cosus, P. defossis, and P. hassleri, but not in specimens of
P. agassizu. No account of the internal genital organs is given
by Faxon.
The best and most recent account of the supernumerary aper-
tures and ducts of Parastacus is by Lénnberg, who describes both
sexes, which differ not only in their internal but also in their
external characters. The species described is P. hassleri. Not
only do the males have supernumerary apertures and ducts in
the 11th somite, but the females also in the 13th somite, in the
position of the normal apertures of male crayfish. Although the
supernumerary ducts have a lumen they are not functional, since
the additional genital orifices in the male are only shallow grooves
and those in the female are closed bya membrane. The additional
openings in the male are closely similar in appearance to the
functional openings of the female. Loénnberg states that he has
found bodies resembling eggs in the testis, but he thinks it im-
probable that they “ can be fully developed, still less of propagative
use.” He draws the conclusion that “in Parastacus hassleri a
partial hermaphroditism is prevailing.” It is interesting to note
that the apertures in most species of Parastacus* are on the same
somites as in the abnormal Astacus described by Benham.
In view of the frequent occurrence of genital apertures in
Nephrops on the basal joints of other legs than the third and
fifth, the coexistence of apertures upon these legs cannot be
regarded as conclusive evidence of a partially hermaphrodite
condition as some authors have supposed. Apertures on the
fourth pair of legs have not, so far as I know, been recorded for
Parastacus, but it is not unreasonable to suppose that if a large
number of specimens were examined they would be found to
occasionally occur.
To those who will regard the abnormal genital openings in
Nephrops as evidence that the apertures and ducts were meta-
merically repeated in past times, all the above-cited cases must be
interesting in view of Lankester’s suggestion that the genital
ducts of Arthropods are derived from nephridia. Allen has
described the genital’ ducts in young adults of Palemonetes as
1 Mr. Borradaile has called my attention to the fact that in male specimens of
Pagurus deformis M.-Edw. the female apertures also normally occur. Vide Borra-
daile, “ On some Crustaceans from the South Pacific, Part IT.,” P. Z.S. 1898, where
references are given.
1902. | VARIATION IN THE NORWAY LOBSTER. 11
agreeing ‘in all their relations with those of Peripatus,” and the
probability of their being derived from nephridia he regards as
“very great.” It is, however, very doubtful whether the case of
Nephrops has any real bearing on this question, seeing that in
Astacus we may also get variation in the direction of reduction of
apertures, and in Cambarus simple homeotic shifting of the
apertures without any addition to their number.
However this may be, the occurrence of such a high percentage
of a particular kind of abnormality as I have recorded among the
Norway lobsters of the Forth area during the present year is
instructive as supplying another example of the falsity of the
doctrine that a well-marked variation cannot exist with any con-
siderable degree of frequency owing to the so-called “swamping
effects of intercrossing.”
Specimens illustrating the various arrangements of the genital
apertures described in this paper were exhibited before the
Zoological Section of the Meeting of the British Association at
Glasgow.
In conclusion I must express my indebtedness to Mr. Bateson,
by whom I was induced to undertake this investigation.
References to Literature.
AtLten.—‘‘ Nephridia and Body-cavity of some Decapod Crustacea.”
Q. J. M.S. vol. xxxiv. p. 403 (1893).
Batrson.—“ Materials for the Study of Variation.” London,
1894.
Benuam.—“ Note on a Couple of Abnormalities.” Ann. & Mag.
Nat. Hist. vol. vii. p. 256 (1891).
BeRGENDAL.—“‘ Ueber abnorme Formen der ersten abdominalen
Anhiinge bei einigen Krebsweibchen,” Bihang till K. Sv.
Vetenskaps-Akad. Handlingar, vol. xiv. Stockholm, 1889;
and ‘‘Neue Beobachtungen iiber die Formvariation der
ersten abdominalen Anhinge bei Krebsweibchen,” 7bid.
vol. xv. 1890.
Conz.—‘ Some Variations in the Spinal Nerves of the Frog.”
Trans. Liverpool Biol. Soc. vol. xv. p. 114 (1901).
DesmArest.—‘“ Note sur une Disposition anormale des Organes
génitaux observée dans VAstacus fluviatilis Fabricius.”
Annales de la Société Entomologique de France, 2° série,
vol. vi. p. 479 (1848).
Faxon.—“ On some Crustacean Deformities.” Bull. Mus. Comp.
Zool. vol. viii. p. 257 (1881).
Faxon.—“ Revision of the Astacide.” Mem. Mus. Comp. Zool.
Camb., Mass. vol. x. 1885.
Faxon.—‘ Observations on the Astacide in U.S. National Mus.
and in Mus. Comp. Zool.,” &e. Proc. U.S. Nat. Mus. vol. xx.
p-. 643 (1898).
Huxtey.—‘‘On the Classification and the Distribution of the
Crayfishes.” P. Z.8. 1878, p. 752.
12 DR. E, LONNBERG ON DIGESTIVE [Jan. 14,
Von Inrrtne.— Parastacus.” Congrés International de Zoologie
& Moscou, Aug. 1892 (dated Rio Grande del Sol, 1892).
Lankester.—‘ Note on Gulland’s memoir, entitled ‘ Evidence in
favour of the view that the Coxal Gland of Limulus and of
other Arachnids is a modified Nephridium.’” Q. J. M.S.
vol. xxxiv. p. 427 (18938).
Lonnperc.— Some Biological and Anatomical Facts concerning
Parastacus.” Zool. Anz. vol. xxi. p. 334 (1898).
Von Marrens.—Sitzungs-Berichte der Gesellschaft naturforsch-
ender Freunde zu Berlin, 1870.
Sr. Grorce.— Ueber eine Zwitterbildung beim Flusskrebs.”
Arch. f. mikr. Anat. vol. xxxix. p. 504 (1892).
2. On some remarkable Digestive Adaptations in Diprotodont
Marsupials. By Dr. Einar Lonnsere, C.M.ZS.
[Received November 18, 1901. ]
(Text-figures 4-6.)
While dissecting for other purposes some Phalangerids, my
attention was attracted by the great difference in the develop-
ment of the intestine in the different species. As some of the
observations made at the time are of a certain interest, the
following account of the comparison of the conditions found in
the different animals may perhaps be acceptable.
Before I proceed to report upon my own investigations, some
preliminary remarks may be made concerning the views of
other authors in similar cases.
The correspondence between an animal’s diet and the develop-
ment of the different parts of its intestine is a well-known fact ;
but, on the other hand, the reason why this must be so has been
comparatively little discussed. Hllenberger, for instance, has
stated that the great development of the cecum in the Horse
stands in connection with its diet, which chiefly consists of matter
rich in cellulose. The food passes in this animal rather rapidly
through the stomach and the small intestine, but is then retained
in the cecum, where, to a great extent, digestion and absorption
take place. In his papers on Rodents, especially in his great
work ‘ Ueber das System der Nagetiere,’ Tullberg has expressed
the opinion that digestion and absorption of cellulose take place
in the cecum and the colon. He says also that the digestion
of this kind of food is not only dependent on the length and width
of these intestinal tracts, but also on the slowness with which the
food passes through these parts of the intestine. There are in fact
to be found many structural adaptations for the purpose of
retaining the food or retarding its passage. The same author
also discusses the reason why some animals among the Rodents,
viz. the Myoxids, have lost their cecum. He believes that such
a reduction is the result of a diet chiefly consisting of such
1902. ] ADAPTATIONS IN DIPROTODONT MARSUPIALS. 13
substances (amylum, fat, proteine), the digestion of which may
take place in the small intestine without the help of any cecum
and without any specialization of the colon. Vegetable food may
thus, just as well as a carnivorous diet, lead to the loss of the
cecum. He also draws attention to the condition found in
the peculiar Phalangerid Tarsipes, in which the cecum is
entirely wanting, and thinks that this depends upon the fact that
this animal feeds chiefly on honey. This last statement is of
special interest because Tarsipes belongs to the same family as
the animals which are to be considered here; and it might, with
regard to the development of its intestine, be put at one end of
the series described below.
The chief material for this little study was afforded by some
specimens of Phalanger maculatus and Petawrus breviceps, collected
in New Guinea by the late Dr. E. Nyman; a specimen of
Pseudochirus occidentalis brought home from Western Australia
by the late Captain Forsstrém; and a specimen of T'richosurus
vulpecula from an unknown locality. In addition to these I have,
with the kind permission of my friend Professor T. Tullberg, had
the opportunity of using other available material in the Zoological
Museum of the Royal University at Upsala, and I beg to offer
him my best thanks for these new proofs of his never-ceasing
liberality.
My first attempt was to try to find out on what kind of diet
the above-mentioned animals lived, by carefully examining the
contents of the stomach and the intestine. The stomach of the
Petaurus contained pieces of the chitinous integument of various
insects and larve, some whole Podurids, and hair of the animal
itself. Among the Podurids my friend E. Wahlgren was able to
distinguish specimens of Jsotoma palustris and of an Achorutes.
It seems accordingly to be certain that this animal may be termed
entomophagous, although perhaps also berries etc. may enter into
its diet. The stomach and the intestine of the Phalanger
maculatus were completely filled with fruit-pulp, and there is
thus reason to regard this Cuscus as chiefly carpophagous. The
stomach in my specimens of Pseudochirus and Trichosurus was
empty ; the contents of the intestine and especially the cecum
indicated, however, a vegetable origin. In the cecum of Zricho-
surus comparatively large pieces of the fibrous skeleton of leaves
could be found, but the parenchymatous substance was digested
or, at any rate, loosened from the “nerves.” This agrees well
with lLydekker’s words—‘“‘the highly aromatic leaves of the
Peppermint-gum form the favourite food of these animals.” !
The cecum of Psewdochirus was filled with a substance in which,
under the microscope, various parts of leaves, upper and lower
epiderm, bundles of vessels, etc. could be discerned. There was
also a good deal of fine sand, which, probably as dust, had once
covered the leaves and sprouts on which the Pseudochirus had
1 LLydekker : “A Handbook to the Marsupialia and Monotremata.’ MLondon, 1894.
14 DR, E. LONNBERG ON DIGESTIVE [Jan. 14;
fed, and thus been swallowed together with the vegetable matter.
Trichosurus and Pseudochirus are thus chiefly phyllophagous,
and so also is the Koala (Phascolarctos), which feeds on Eucalyptus-
leaves. Of the latter I have had only scant material—the dried
cecum of a grown animal and the intestine of a foetus. It may,
however, also be considered in this comparison because its anatomy
is known from the descriptions of Owen * and Forbes’.
For the purpose of illustrating the length of the different parts
of the intestine, the measurements are given in the following
table * :—
Trichosurus. Pseudochirus. Phalanger. Petaurus.
Length of animal
without tail ... 39 cm. 28°5 cm. 58 cm. 14 cm.
Small intestine... 213 ,, 139°5 ,, 198 ,, 48 ,,
@xcumye ee 23 ,, 42°5 ,, 69 ,, 5'5 ,,
Large intestine ... 122 ,, 87-2 ,, 261 ,, Des
To make the comparison easier it is, however, convenient to
express the relation between the length of the different parts of
the intestine and the length of the animal itself (without tail).
This is donein the following table, in which the numbers indicate
percentages of the length of the animal without tail. The
numbers under the head of Phascolarctos are calculated from the
measurements of this animal recorded by Forbes (J. c. p. 184).
There are also added measurements taken by myself from a
specimen of the small insectivorous Acrobates pygmeus.
In percentage of the
animal’s length Phasco- Tricho- Pseudo- d Acro-
without tail. Jlarctos. surus. chirus. Phalanger-“Eetavrus: bates.
Small intestine... 642 546 496 360 342 250
Crceum ............ 321 59 149 125 39 22°8
Large intestine... 784 312 305 474, 78 (mutilated)
A glance at this table reveals that in the Koala all parts of
the intestine are very much more lengthened than the corre-
sponding parts of the intestine of the animals at the other end of
the series. The cecum and the large intestine are considerably
larger even than in the likewise chiefly phyllophagous Trichosurus
and Pseudochirus. On the other hand, all three phyllophagous
animals have the small intestine developed in comparatively the
same degree but a good deal longer than in the others. The
reason why the cecum and the large intestine in Jrichosurus
and Pseudochirus are comparatively shorter than the same organs
in the Koala may be seen from the description further on. If
the digestion of the cellulose takes place chiefly in the cecum,
as has been supposed by the authors quoted above and with
1 « Anatomy of Vertebrates.’
2 “On some Points in the Anatomy of the Koala (Phascolarctos cinereus).”
P. Z.S. 1881, p. 180.
3 All measurements are taken by means of a thread laid along the middle of the
intestine while adherent to the mesentery.
1902. | ADAPTATIONS IN DIPROTODONT MARSUPIALS, 15
whom I fully agree, it may be asked—why has, then, the small
intestine become lengthened in the three phyllophagous animals 2
I think that this may be explained in the following way :—The
leaves of which the food of the animals is composed consist not
only of cellulose, but contain also protoplasmic, amylaceous, and
other substances, which ought to be digested and re-absorbed in
the small intestine. These substances are, however, all of them
enclosed within the membranes of the cellules of the leaves, and
these membranes are more resistant than, for instance, the thin-
walled cellules composing the pulp of fruit. The food derived
from leaves must consequently be subjected to a longer treatment
also in the small intestine, before yielding its useful substances,
than food consisting of fruits needs—not to speak of animal food.
The small intestine of Trichosurus is villous, as has already
been remarked by Oppel’, but my material does not allow any
description of the villi. About 93 cm. from the opening into the
large intestine I have found two roundish Peyer’s patches,
situated near each other and measuring respectively 3 and
23 mm. in diameter. They are solid and not composed of small
nodules. In the intestine of the Koala there are no Peyer’s
patches according to Forbes (/. c. p. 184). The last portion of
the ileum is in 7’richosurus conspicuously more thick-walled than
other parts of the small intestine. Its mucous membrane forms
distinct longitudinal plice, and it seems thus to be more rich in
glands than other parts. In addition to this there are to be
seen what I am inclined to term, with Owen, some “ wide and
deep glandular fosse.” The largest of these is situated about
1 em. from the ileo-cecal opening, and measures nearly 5 mm. in
length by 13 in width. About 1 cm. higher up the ileum there
is another one of the same kind although smaller, so that it
measures only 2 mm. in length. There are also indications of
some other depressions, but they are shallower and less distinct.
My material does not allow of any histological investigations, but
I hardly think I can be much wrong in interpreting these as
accumulations of glands. The ‘“fossee” mentioned by Owen in
the words quoted above were found by him in the large intestine
of the Koala, and are thus not homologous with these. The
“‘fosse ” found by Owen have, however, their homologue in a
thickened glandular area, with numerous shallow depressions,
situated on the adjoining borders of the colon and the cecum
of Trichosurus just opposite the ileo-ceecal opening. The ileo-
cecal valve is well developed and protrudes into the colon. The
limit between the colon and the cecum is only marked by a short
plica from the ileo-cecal valve and a weak sphincter ceco-colicus.
If this sphincter is weak it is assisted in its functions by a series
of cecal sphincters which are strongly developed. Their number
is four. The first is situated about 3 cm. from the czco-colic
one. The next is stronger and found at about the same
1 ‘Lehrbuch d. vergl. mikroskopischen Anatomie der Wirbeltiere, Zweiter
Teil (Jena, 1897), p. 288,
16 DR. E. LONNBERG ON DIGESTIVE [Jan. 14,
distance from the first. The third, which is almost the strongest
lies still 4 em. nearer the blind end. The fourth, which is about
equal to the third in strength, is situated at a distance of about
8 cm. from the tip of the distal end. These two last-mentioned
sphincters are 33 to 4 mm. thick, and protrude in the preserved
state, as circular valves 3 mm. or more, into the lumen of the
cecum ; and there is no doubt that in the living animal they are
capable of entirely shutting off one portion of the cecum from
the other, thus retaining the enclosed food during a suitable time
Text-fig. 4,
Cecum of Trichosurus vulpecula. Nat. size.
for decomposition. The walls of the caecum increase considerably
in thickness towards its blind end; and it is evident that this
increase includes the muscular coat as well as, and that especially,
the glandular layer. This is the reason why the sphincters also
must have an increased size and strength towards the blind end.
The mucous membrane of the cecum is, at least from the third
sphincter and onwards, transversely plicated, the plice becoming
more prominent towards the blind end. They do not extend,
however, as simple plice all round the cecum, but the ridges
anastomose now and then so that they form a network with
1902, ] ADAPTATIONS IN DIPROTODONT MARSUPIALS. 17
transversely very much elongated, but longitudinally quite
narrow, meshes.
The figure (text-fig. 4) represents the blind end of the cecum
from the last sphincter, and shows also the transverse plicee
described above.
When the cecum is filled the blind end seems to taper toa
point not unlike a vermiform appendix.
The colon is thin-walled and smooth, only showing, in places
where it is contracted, longitudinal folds, which become effaced
by stretching.
From this description it will be evident that the cecum of
Trichosurus is an organ which has become to a considerable
degree specialized for digestion (and re-absorption). When the
chyme passes through the terminal portion of the ileum it
becomes mixed with the secretion of the glands of that intestinal
tract, to which is added, when it enters through the ileo-cexcal
valve, the secretion from the czco-colic glandular patch. Thanks
to the sphincters and the well-developed muscular coat of the
cecum, the food can be moved backwards and forwards, or retained
in the cecal divisions, and then be subjected to the action of the
glands through the development of an increased surface due to
the transverse plice. This great specialization gives a satisfactory
explanation why the cecum of TZrichosurus does not need to
grow out to such a size and attain such a capacity as that of the
Koala, It might be questioned whether any proofs can be given
to show that cellulose is really decomposed and digested in the
cecum of this animal. It may then, firstly, be referred to the
peculiar structural specialization described above; secondly, it
may be stated that the contents of the colon which have passed
the cecum seem to indicate such a digestion. There may still be
recognized remains of the most resistant parts of the vegetable
tissue, such as pieces of epiderm, isolated sclerenchym-cellules,
bundles of vessels, ete.—all of them looking as if they had been
cleaned by some reagent, so that only the hardest ‘“ skeletal”
parts were left. There are also to be seen the spiral threads,
these being the only remains of spiral vessels, the thin walls of
which probably have been digested. But I could not detect any
parenchymatous cellules or other softer parts. I think, there-
fore, that it may be admitted that the softer cellulose has been
decomposed and digested, leaving only the more or less lignin-like
substance.
In Pseudochirus the small intestine is, as usual, thin-walled
and villous but otherwise smooth. The walls of the ileum do not
seem to show any increase in thickness, The ileo-czecal valve (v.1.¢.)
is well-developed, and from it extends as a transversal fold a
execo-colic valve (v.c.c.) and sphincter, as may be seen in the figure
(text-fig. 5, p. 18).
Close by, but on the colic side and also near the ileo-cxcal
valve, a brownish glandular patch is seen, homologous with that
described above and in a similar situation in 7’richosurus. (It is
Proc, Zoot, Soc,—1902, Vor. I, No, II, 2
18 DR. E, LONNBERG ON DIGESTIVE [Jan. 14,
not represented in the figure.) The cecum of Pseudochirus is a
good deal larger than that of Trichosurus, as the above-recorded
comparative measurements indicate, and offers also quite another
aspect (compare figures). It is provided with two very strong
teenie, which continue to the blind end of the cecum. These
tenie have an average breadth of 3 mm. and extend one on
each side. By means of these teenie and the mesentery the wall
of the cecum is folded so that it forms three series of sacculi.
Text-fig. 5.
\ : :
Rirawen ee
Cexcum ot Pseudochirus occidentalis. Nat. size.
c., colon; 7., ileum; v.¢.c., cxeco-colic valve; v.7.c., ileo-czecal valve.
Through this the effectiveness of the cecum as a digesting organ
is greatly increased, the more so as the depth of the pockets is
comparatively great. The end of the cecum in this animal
is bluntly rounded, and thus different from that of Trichosurus.
The first part of the colon is longitudinally plicated, but the
plice are not strongly developed and soon disappear.
1902. ] ADAPTATIONS IN DIPROTODONT MARSUPTALS, 19
The digestion in the cecum seems to be rather complete, since
at a distance of 25 cm., more or less, from the ileo-ceeal valve the
fecal matter is already formed into balls. A microscopical
investigation of these fecal remains shows that they are chiefly
composed of pieces of thick-walled epiderm, bundles of vessels,
isolated prosenchyme-cells, and similar matter. But the softer
vegetable tissue has disappeared and the spiral threads of the
vessels are isolated, indicating a digestion of the substance that
once formed the walls.
The intestine of the Koala has been described, as already
mentioned, by the authors quoted above. Only a few remarks
will therefore be made here concerning the intestine of a
marsupial foetus of this species measuring about 9 cm. in length.
Its small intestine measured about 37 cm., the cecum 8°5 cm., and
the large intestine about 29 cm. The length of the three different
parts of the intestine, compared with the length of the fcetus
itself, is thus expressed by the following percentages: 411, 94,
322. If these now are compared with the corresponding ones
from a grown animal, calculated from Forbes’s measurements
(see above), the difference is quite striking with regard to the
cecum and the large intestine. The former is proportionately
only about a third as long in the foetus as in the full-grown
animal, and the latter less than half as large in the feetus as in
the adult. The difference of the small intestine of both stages is
not so great, that of the fetus being about four-fifths of the same
in the adult. It is also to be remarked that in the feetus the
small intestine is considerably longer than the colon, but in the
adult the reverse condition prevails. These differences can of
course be ascribed to the difference of the diet of both stages.
The milk food of the fcetus is chiefly or completely digested in
the small intestine, but the vegetable diet of the adult needs a
greatly developed cecum and colon. The longitudinal folds of
the cecum and the colon are, however, already developed in the
foetus.
The condition found in the Wombat is very peculiar, The
narrow opening of the ileum protrudes, surrounded by an “ ileo-
cecal” valve, into the colic cavity. This valve has very broad
lips, and within the same opens the lumen of the “ processus
vermiformis” (Owen), only separated from the opening of the
ileum by a septum—that is, in other words, the terminal portions
of the vermiform appendage and of the ileum are thus fused
together into one structure protruding into the colon ; both open
with separate orifices, which are, however, surrounded by the lips
of the same valve. The glandular patch described above as
situated near the valvula cceco-colica in the related Phalangerids
is found in this animal too, but extends partly on to the outer
side of the “ ileo-ceecal” valve itself already mentioned. From
this latter valve plice extend on two sides. These plice seem to
stand in the same kind of connection to the ileo-czecal valve as in
normal cases the czeco-colic valve does, But they do not extend
Q*
20 DR. E, LONNBERG ON DIGESTIVE [Jan. 14,
transversely in the colon as the latter is apt to do, but longi-
tudinally, so that one plica extends down the colon, the other in
an opposite direction on the other side. J am, however, inclined
to think that these plice morphologically correspond to the more
or less developed cxco-colic valve of other related forms. Although
the direction is different the connection with the ileo-czcal valve
may decide in favour of such an interpretation. These plice do
not form any boundary between the cecum and the colon. They
form no boundary at all directed as they are now. But how
could they get such a direction? I think this may be explained
im connection with the shape of the cecum. Owen, in describing
the intestine of the Wombat, used the following words :—“ The
cecum is extremely short but wide; it is remarkable for being
provided with a vermiform appendage.”’ Later authors have
adopted this same interpretation, but I hardly think it is right.
If it had been a true vermiform appendage, that is, the reduced
blind end of a cecum, it ought to have opened into the cecum of
which it itself was a part. But it does not, as has been already
stated above. It opens with a quite independent opening of its
own near that of the ileum. I judge from this that the so-called
processus vermiformis of the Wombat represents a rudiment of
the whole cecum. If we assume that a moderately developed
cecum should for some reason or another become reduced toa
mere appendage, it must acquire a similar situation and open into
the colon close to the ileum. It might then easily happen that
the wall of the terminal portion of the rudimentary cecum
became fused with the wall of the ileo-cecal valve. Such an
event might be the more easily efiected as the shortened
mesentery of the cecal rudiment would draw the latter more and
more to the ileum and make both more closely connected. It
would also be more convenient if the two openings into the colon
lay near each other and were parallel in direction, because there
would then be less risk of particles of food entering the cecal
rudiment. I believe, indeed, that such a retrograde development
has really taken place, and that in the ancestors of the Wombat
the cecum has been reduced to a rudiment, which might happen
if they lived on such a diet that the cecum was not needed for
the digestion of the food. When the cecum had already reached
a considerable degree of reduction, the diet of the animals was
changed, and they began by-and-bye to feed on harder and legs
easily digestible vegetable matter containing cellulose, ete. The
czcum was now, however, so rudimentary that it could not, as in
other related forms which live on a similar diet, help in the
digestion of this food-stuff. This function became, therefore, the
duty of the colon alone, which in consequence had to be con-
siderably enlarged. It grew in strength, and its capacity increased
so that it would be able to hold the greatly augmented amount of
the less nourishing food that was needed for the sustaining of life
and growth. Thecolon was then distended by the large quantities
1 Owen: ‘ Anat. of Vertebr.’ p. 417.
1902. ] ADAPTATIONS IN DIPROTODONT MARSUPIALS, FAL
of food-material, and the mechanical pressure of this heavy load
might have produced expansions, which, if localized, formed bags
or sacculi. The first part of the colon might be expected to have
been strongly affected by this pressure. It is consequently
natural that some wide sacculi should be formed there, and it is
these distensions which have been described and figured by the
authors as cecum, although they are derived purely from the
colon. It is also easy to understand that when this distension
took place the originally transverse czeco-colic plica was drawn or
turned out of place to its present longitudinally-running direction.
That the mechanical pressure of the contents of the colon
has really played important parts in transforming it to its
present shape, may also be proved by another fact. By a broad
band opposite the mesentery the colon is sacculated, which, of
course, is also an adaptation to its function. At the place where
the colon is most closely, by a very short mesentery, soldered to
the back of the abdominal cavity, the pressure of the contents
would, thanks to this fixation, be more effective. There has thus
been formed two large sacculi, which give the colon at that place
a size amounting to twice that of its usual width. The shape and
size of these sacculi are identical in two specimens which I have
seen. Thisconfirms the correctness of the statement ; and I think
it is these which Owen means when he says: ‘One of these
sacculi was so much longer than the rest as to almost merit
special notice as a second cecum.”
Peyer’s patches of comparatively large size, 1 to 2 cm. in
diameter, are scattered in considerable numbers in the walls of
the colon, especially in its middle parts.
The material which I have used for this study has long been pre-
served in spirit, and the measurements are perhaps therefore not
somuch tobe relied upon. It may, however, be mentioned that the
small intestine measured in one specimen about 410 cm., the cecal
rudiment 6 cm. from its blind end to its opening, and the large
intestine 840 cm. The “secondary cecum” is situated nearly at
the middle, or 430 cm. from the end, Even if these measure-
ments are imperfect in the detailed statements, they show satis-
factorily that the large intestine has been strongly developed.
Probably it is fully eight times the length of the animal, or even
more than in the Koala.
The interior surface of the duodenum in Phalanger shows
very plainly a reticulate structure, larger primary and smaller
secondary plice may easily be distinguished. It offers thus some
faint resemblance in appearance to the structure of the reticulum
of a ruminant. The plice are in both cases formed by coalescence
of papille. The villi of the intestine are well developed on the
ridges forming the network, but some are also scattered in the
interspaces. Lower down the small intestine this reticulate
structure is less conspicuous, but my material is not in such good
condition that I can say where it entirely disappears. The
jejunum appears, however, quite smooth. it
22 DR. E. LONNBERG ON DIGESTIVE [Jan. 14,
As in Zvrichosurus, the small intestine of Phalanger is pro-
vided with at least one Peyer’s patch. It is in the specimen
before me situated 74 cm. from the cecum, and measures 30 mm.
in length by 13 in breadth, being composed of a great number of
small nodules. There are, however, probably more than one
Peyer’s patch normally in the small intestine of the Cuscus, since
Cunningham observed no less than nine in his specimen described
in the ‘Challenger’ report. Some of these were, however, “a
mere speck.” ‘The terminal portion of the small intestine shows
some longitudinal folds, but these are probably not permanent as
they disappear by transverse stretching. The ileo-czxcal valve is
well developed and protrudes 12 to 14mm. into the cecum. From
this valve extends on both sides a fold—the ceeco-colic valve. At
the ileo-cecal valve it is about 7 mm. in height, but gradually
diminishes; about 1 cm. from the valve it passes into the
muscular thickening which forms the cxeco-colic sphincter. The
communication between the cecum and the colon may thus be
completely shut off by means of the incomplete ceco-colic valve
and by contraction of the ceco-colic sphincter. When such a
shutting-off is effected it seems as if the opening of the ileo-cecal
valve would be directed into the cecum, and the function of the
above-described caco-colic valve may partly be to brace the ileo-
cecal valve so that it shall not be compressed and closed when the
ceeco-colic sphincter contracts. But, as it is arranged now, the
contents of the small intestine may pass directly into the cecum
without risk of slipping down into the colon. On the cecal side
of the valve there is an area on which the mucous membrane is
provided with a considerable number of small depressions. These
are about 1 mm. in diameter, and correspond, no doubt, to the
glandular patch with similar depressions which has been described
above in the phyllophagous Phalangerids, although the situation
is a little different in these latter,in which this patch is found on
the colic side of the valve. Cunningham does not mention this
glandular patch in his description of the intestine of the Cuscus.
The width of the cecum is different at different places. It is
at first about 4 cm., then widens to 6 cm., but soon becomes
constricted to only 24 cm., widens again to 53 cm., then it 1s
constricted to 24 cm,, and again widened to 4 c¢m., which con-
dition is once more repeated, and then it finally tapers towards
the end, which terminates in a digitiform appendix 2 em. in length
by 4mm. in thickness. Cunningham* has in the same species
only observed that the cecum “tapers uniformly.” ‘The appendix
is hollow and filled. with the contents of the cecum. Its walls are
thicker than those of the ordinary cecum, and it might be a
lymphatic organ, which perhaps might be compared with the one
of similar situation in the common rabbit.
The cecum of Phalanger is somewhat sacculated by means of
J) B)
1 “ Report on the Marsupialia,’” Rep. Scient. Results ‘Challenger,’ Zoology, pt. xvi.
p. 161.
1902. | ADAPTATIONS IN DIPROTODONT- MARSUPIALS. 23
mesenteric bands of muscular fibres which are most often longi-
tudinally, but sometimes obliquely, arranged. Such bands are
found on both sides of the cecum. The interior of the cecum
shows at the constricted places slight longitudinal folds, which,
however, probably are of a temporary nature. They are thus
not to be compared with the longitudinal folds described in the
Koala by Owen and Forbes (J. ¢.).
In the large intestine of Phalanger there are some longitudinal
folds near the upper sphincter. They are, however, quite short
and continue but a few centimetres from the ileo-cecal valve,
and are therefore quite unlike the longitudinal “ valvule conini-
ventes” described in the Koala by the authors just mentioned.
The colon tapers abruptly from the width of the cecum 4} cm.
to 23 and then to 13cm. In places where it is much distended
its width may reach 3 cm., but as a ule it is less than that of
the small intestine, usually 2-14 cm.! The rectum attains a
width of 23 cm., and is provided with about half a dozen longi-
tudinal folds, plainly conspicuous, but not much developed.
They may, however, be regarded as homologous with those of
the Koala in a corresponding situation and of nearly the same
number, according to Forbes. The non-digestible remains in
the colon of Phalanger consist of pieces of epiderm of fruit, pro-
senchymatous fibres and vessels. The main mass of the fruit-
pulp is, however, so decomposed that its particles cannot be
identified.
The duodenum of Petawrus is very densely beset with flattened
more or less tongue-like villi which are transversely arranged
and partly form thin denticulated lamelle. They lie so close
together that the contents of the intestine probably only, or
at least chiefly, come into contact with the tips of the villi—
a condition found by Oppel (J. ¢. pp. 288-9) in Zrichosurus.
In Petaurus the small intestine is beset with villi through its
whole extent, although they decrease in size posteriorly. The
condition found in Acrobates seems to be essentially the same.
In Petaurus the duodenum forms a much more distinct loop than
in Phalanger ; it is 3 em. in length, the ascending branch being
closely connected with the descending one and returning along
the same to the pyloric tract. In the latter the duodenal loop is
less pronounced because the ascending branch is only half as long
as the descending one.
In Petawrus the connection between the large and small intes-
tine takes place in such a manner that the ileum opens indo the
colon, into which the valvula ileo-cwcalis (the name is thus not
quite suitable in this case, more correctly v. ileo-colica) protruded
about 2mm. Close to this valve there is between the cecum and
the colon a strong sphincter, partly like a valve protruding into
the cavity of the colon and only leaving a very narrow opening
(which, of course, also can be closed) between the same and that
of the cecum.
1 All such measurements are taken across the empty but not opened intestine,
24 DR, E, LONNBERG ON DIGESTIVE [Jan, 14,
It is thus evident that the contents of the ileum must pass
into the colon, and from there, when the sphincter mentioned
above relaxes, into the cecum. It is also evident that no large
pleces can pass through the narrow opening into the cecum. A
comparison between the contents of the colon proves this state-
ment completely. A sample of the contents of the colon taken
2 cm. from the ileo-cecal valve consists chiefly of large pieces of
the chitinous integument of insects, sete of such animals, ete.
A sample from the cecum consists only of the tiniest particles
which cannot be measured or identified. The narrow opening
between the cecum and the colon serves accordingly as a filter.
The large indigestible remains are kept back in the colon, the
fluid and the fine particles suspended in the same pass into the
cecum, where, no doubt, an absorption of the fluid takes place,
after which the indigestible remains are forced back to the colon
to be expelled with the feces. The function of the cecum may
thus be termed absorbing and desiccating. The large intestine
acts nearly in its whole extent as a rectum, as the fecal matter
is already formed into balls at a distance of only 2 cm. from the
ileum (text-fig. 6),
Text-fig. 6.
Cecum of Petawrus breviceps.
ec, cecum; 7, ileum; co, colon. Valvula ileo-colica is seen to protrude into the
colon, and a bristle is inserted through the narrow opening of the cxco-colic
sphincter.
In Phalanger, on the other hand, it can be assumed with
certainty that the cecum has a digestive function, which may be
concluded from the fact that it 1s provided with large glands.
The great length of the colon makes it probable that it has
digestive as well as reabsorbing powers.
The dimensions of the different tracts of the intestine in a
marsupial young of Petawrus breviceps, measuring 63 mm. in total
length without tail, were as follows:—Small intestine 185 mm.,
cecum 17 mm., and large intestine 43 mm. If these measure-
ments be compared with those of the young animal itself (without
tail) the relation is expressed by the following percentages :—290,
26, and 68. From this it may be seen that all three parts are
1902.] ADAPTATIONS IN DIPROTODONT MARSUPIALS. 25
somewhat shorter in the young than in the adult. The difference
is, however, not so great as in the case of a marsupial fcetus of
the Koala already mentioned. Although the latter represented
a younger stage’, the conclusion can thus be drawn that the
milk diet of the young Petauwrus differs, with regard to its com-
position and therefrom resulting influence on the intestine, less
than it does in the case of the young Koala. It is also of
interest to note that in the young Petawrus the cecum stands in
quite open communication with the colon. That is because the
sphincter is not needed yet for the purpose of prohibiting any
indigestible remains from entering the cecum as in the adult.
In Acrobates the arrangement of the connection between
the small and the large intestine is the same as in the adult
Petaurus. The ileum opens with its valve protruding like a
mouthpiece into the colon; and there is a very strong constric-
tion between the latter and the cecum. Although the stomach
in my specimen of Acrobates was empty, I think it may be
assumed that it lives on a similar diet to Petawrus; and at any
rate the function of the cecum seems to be perfectly alike in
both animals.
The small intestine of Acrobates is comparatively shorter even
than that of Petawrus. In the latter and in Phalanger the
length of the small intestine is comparatively not much different.
This may be understood as meaning that that part of the food-
material which ought to be digested in the small intestine of
Phalanger is not difficult to digest. It may chiefly consist of the
juice and other contents of the soft parenchymatous cells of the
fruit-pulp. This matter is, of course, more easily accessible for
the digestive organ than is the material contained in the better
protected cells of the leaves etc., which form the food of the
animals considered above, and the small intestine of which, there-
fore, has become lengthened.
_ Ag the last stage in this series, showing a different develop-
ment of the intestine and especially of the cecum in accordance
with the different diet, Zarsipes may be mentioned; this animal
has, as already remarked, entirely lost its cecum, because such an
organ is superfluous for a honey-eater.
The general arrangement and structure of the dentition of
these animals indicate that also with regard to those parts adapt-
ations for different purposes have taken place. The dentition of
Phascolomys is evidently most specialized. Its incisors are more
reduced in number than in the others, the canines are absent,
and the molars have persistent pulps. The latter are also curved
in such a way that the upper molars are laterally concave and
the lower ones are laterally convex. This development and shape
1 The Koala foetus was still naked. The young Petawrus was beginning to
become hairy, so that, for instance, the dark vertebral stripe was well conspicuous,
but the hairs of the tail were not yet prolonged. It had certainly not yet partaken
of any other food than milk, because the mandibular incisors, although protruding
3 mm. from the sockets, had not cut the gum.
26 DR. E, LONNBERG ON DIGESTIVE [Jan. 14,
of the molars reminds one of the same in the Hares and Rabbits.
In his above-quoted work Tullberg has already drawn attention
to this parallelism. The explanation given by that author of
the development of persistent pulps in the molars holds good
for the Wombat as well. He believes that such teeth have
been developed chiefly in animals which feed on hard and tough
roots. Such animals must take much sand into the mouth when
feeding, and the sand must act strongly upon the crowns of the
molars in the act of grinding. This renders persistent growth
necessary. That the food of the Wombat consists mainly of
roots is a well-known fact; anda glance at the crowns of the
molars suffices to show the marks of the sand as transverse
scratches. Although the molars of the Wombat in their general
shape resemble those of a Rabbit, the chewing must take place in
quite a different manner in both forms', as can be seen from a
comparison of the structure of the mandible in both animals.
The dentition of the Phalangerids has been described by Flower
and lydekker in their valuable manual*. They have drawn
attention to the “crescentoid” cusps of the molars in Pseudo-
chirus and in the Koala “recalling those of the Selenodont
Artiodactyle Ungulates.” This ‘“subselenodont” dentition is, of
course, very suitable for phyllophagous animals. It becomes the
more effective because the distance between the upper molar
series is greater than that between the mandibular molar series,
so that the outer row of cusps of the latter fit in between both
rows of the upper. Through this arrangement the jaws get as
it were a cutting-power, and when the lower jaw is moved side-
ways the sharp enamel ridges have a great power of tearing and
grinding the food. It is, in fact, evident that the chewing of the
food takes place in the following manner :—The lower jaw is
moved towards one side so much that the outer margin of its
molars corresponds to the outer margin of the upper molars. If,
then, both jaws are pressed hard against each other the lower
jaw must glide, with triturating effect, ina median direction—
in consequence of the fact that the main surface of the upper
molars slopes inward—till the outer cusps of the lower molars fit
in between both series of cusps of upper molars as described
above. Then the same movements are repeated again on the
same or the other side. The movements of the jaws in the act
of chewing may thus be compared with those in the Ruminants
—as might be expected seeing that the teeth have a similar
structure and position.
The incisors in these two animals are different in shape. In
the Koala they are comparatively narrower than in Pseudochirus.
The compressed mandibular incisors glide with their bevelled
ends inside the upper median incisors, and work against the
second pair of upper incisors; the latter in their styliform shape
z This is also remarked by Tullberg (7. ¢.).
* An Introduction to the Study of Mammals. London, 1891.
1902.] ADAPTATIONS IN DIPROTODONT MARSUPIALS. 27
and situation behind the first pair remind one of those of the
duplicidentate rodents. The incisors in this animal serve thus to
nip off pieces of the leaves. In Pseuwdochirws the lower incisors
are broad and have sharp cutting-edges also on the sides. They
therefore work together with the upper incisors as a pair of
scissors cutting off pieces of plants and leaves. Both halves of the
mandible are in this animal movable, whereby the cutting-power
of the lower incisors becomes more effective. They may thus
be compared in shape and in action with those of the Kangaroos.
In Trichosurus the incisors ave intermediate in shape between
those of the two animals just referred to. They are broader
than in the Koala, but have a cutting-edge only in front. In two
skulls of this animal before me it is plain that the lower incisors,
when used, are able to work against all three pairs of upper
incisors, Which are all worn—the median ones, however, in such
a manner that a sharp edge is left in front. The two halves of
the mandible do not seem to be movable.
The subselenodont type of the molars is not so prominent in
this animal—at least not when the teeth are worn. The shape
and position of the molars seem also to be different in Z'richo-
surus, because, at least in the specimens before me, the surface of
the two anterior upper molars slopes inwards and that of the
two posterior ones outwards. Inthe lower jaw, in correspondence
herewith, the two anterior molars slope outwards and the two
posterior ones inwards. The crown of the posterior premolar in
each jaw slopes in the same direction as the anterior molars of
the same series close to which it is situated. In consequence of
this arrangement, the upper premolar and the anterior upper
molars effect the gliding in a median direction of the lower jaw
when both jaws are pressed against each other in the manner
described above; but the posterior upper molars arrest the lower
jaw and hinder it from gliding further than to its normal
position. In connection herewith is also to be observed that the
mandibular molar series of 7richosurus—thus differing from the
Koala and Pseudochirws—have not a shorter distance inter se along
their whole length than the maxillary molar series. In TZricho-
surus the molar series of both jaws are, posteriorly, almost
opposite each other, and only anteriorly have the mandibular
molars a more median position than the upper molars. This
accounts for the different direction of the anterior and posterior
molars.
The teeth of a young Phalanger differ a great deal from those
of the old one of the same species. In the half-grown animal
the lower incisors appear to be absolutely broader than in the
adult.- This is, however, only apparently the case. The breadth
is about the same in both. The incisors of the young are thus
not only comparatively broader, but their shape is also different.
They are much more flattened than in the adult and have sharp
lateral edges, so that they resemble the corresponding teeth of
Pseudochirus described above, or of a Kangaroo. The resem-
28 DR. E, LONNBERG ON DIGESTIVE [Jan. 14,
blance is the greater because they have a more horizontal direction
than in the adult, in which latter they are also stouter, com-
pressed, and only provided with an edge at the end. In
the young the lower incisors, on account of their shape,
work against the two median pairs of the upper incisors when
the jaws are shut. In the adult they work only against the
inside of the median pair of upper incisors when in a normal
situation close to each other. The halves of the lower jaw are,
however, movable, more so in the young than in the adult. This,
together with the sideway movements of the lower jaw, explains
also why the second pair of upper incisors are worn. The faculty
of moving the mandibular halves so that the lower incisors may
be separated from each other in the act of biting is, of course,
very useful in many cases’. Thus, for instance, the animal is
capable of securing a much larger piece of some soft fruit ° in
one bite through this arrangement, and when occasionally preying
upon animals or birds this faculty is also of importance. The
mobility of the mandibular halves consequently serves here other
purposes than in the Kangaroos and Pseudochirus.
The upper canines are well developed in young and adult. The
molars of the young Cuscus show four well-developed pyramidal
cusps with radiating ridges, so that, as has been shown by
O. Thomas, they resemble in some degree those of the Koala.
The enamel of these cusps is, however, less developed in the
Cuscus, so that they are in the adult animal soon worn down to
such an extent that the crown becomes almost even, and only
peripherally surrounded by enamel. The teeth are then not
much adapted for any grinding action. The situation and
different sloping of the posterior and anterior molars are similar
to those described in Zrichosurus. The action of the jaws
must consequently be similar, although the enamel is rather less
developed. To crush the pulp of fruits and similar matter the
teeth are, however, sufficient. The hindmost premolar of both
jaws lying just in front of the molar series is somewhat more
strongly developed than in Z’richoswrus, pointed and reminding
one a little of a canine. Those of the upper and lower jaw do
not touch each other as in 7richosurus, but the mandibular
premolar goes inside and in front of that of the maxillary. This
development of the last premolar may have some connection with
the alleged occasionally predatory habits of the animal, —
In Petauwrus the median lower incisors are very long and
slender, The median pair of upper incisors are longer than the
others. They may thus, together with the lower incisors, form a
pair of pincers. It is also probable that the mandibular incisors
themselves may, because both halves of the lower jaw are quite
movable, act as a suitable implement for pinching and scratching
1 The mobility of the mandibular halves of the Rodents and its causes have been
extensively discussed by Tullberg in his work quoted above, p. 345 and following.
2 Tullberg has stated that ‘Squirrels feeding on mushrooms separate their
incisors (J. c. c.).
1902. ] ADAPTATIONS IN DIPROTODONT MARSUPIALS. 29
small insects from their refuge in flowers, in cracks in the bark,
and similar places where these slender incisors may conveniently
be inserted. When securing larger insects this can, of course,
be more easily done with the incisors separated so that they act
as a fork, than if they lie close together and form only one point.
The molars have four moderately developed bluntly pointed
cusps. The lateral row of cusps of the mandibular molars fits in
between both rows of cusps of the upper molars.
On the whole the dentition may be said to approach the
insectivorous type. The molars can certainly not be used for the
grinding of any hard vegetable matter, and the incisors are too
weak to gnaw.
In Acrobates the development has gone still further in the
same direction. The median lower incisors are long and slender,
although, if compared with the skull itself, not so long as in
Petaurus, which has a shorter, less pointed snout. They may
certainly serve as pincers and the mandibular halves are quite
movable. The premolars of Acrobates are much better developed,
longer, and more pointed than those of Petawrus. When the
jaws shut, the premolars of the upper and lower jaws meet, and
the latter slide up in front of the former. These teeth may thus
help in catching and holding the prey, which is not the case in
Petaurus. In the latter the premolars and second incisors of the
lower jaw are small and functionless. This is because, in con-
sequence of the length of the median incisors and the corresponding
shortness of the jaw itself, there is formed a considerable opening
between the upper and the lower jaw corresponding to the canine
and premolar region of the maxillary. The maxillary teeth thus
cannot meet the mandibular teeth, which do not even lie opposite
to them. The molars of Acrobates are similar to those of
Petaurus, but their cusps are sharper. It may be in conse-
quence of the arrangement of the premolars and their use that
Acrobates has been able to reduce its number of molars to 3/3
when Pefauwrus has 4/4.
In none of the Phalangerids which have the rami of the lower
jaw movable, as described above, have I been able to detect in
my material any trace of such a transverse muscle as that which
is found in the Kangaroos at the base of the mandibular incisors,
and which has the function of approximating the inner edges of
these teeth. In the Kangaroos it is said by Leche* that the
mandibular incisors are separated from each other by the com-
bined action of the musculi biventer, mylohyoideus, and genio-
hyoideus. In his great work on the Rodents already quoted,
Tullberg states that m. masseter serves to break or bend the lower
margin of the mandible outwards, and that in such a case the
incisors become pressed close to each other. On the other hand,
the m. transversus mandibule, when contracting, approaches the
lower margins of the mandibular rami towards the median line,
1 Bronn: Kl. u. Ordn. d. Thierr., Satigethiere, vi. 5. 1. p. 681.
30 DR. E, LONNBERG ON DIGESTIVE [Jan, 14,
whereby the incisors become separated. In the Phalangerids in
question I think that in a similar manner as in the Rodents the
masseter may press the mandibular incisors together, but the
pterygoideus intermus separates them from each other. The mylo-
hyoideus seems to be rather weak, and is inserted too high up on
the inner surface of the mandible to have any power of bending
the lower parts of the mandibular rami inwards and thus
separate the incisors. The pterygoideus internus is very strongly
developed, and the angle of the mandible is much inflected so as
to give this muscle a wide area of insertion. It is of interest
to see that in those Phalangerids which have especially movable
mandibular rami, viz. Pseudochirus, Petawrus, and Acrobates,
this inflexion is much stronger than in the Koala with fixed
mandibular rami. This fact gains importance by the obser-
vation that in the three former less is done for the enlarging of
the area of insertion of the masseéer on the outer side of the
mandible than in the Koala, in which the angle is considerably
expanded on the outer side.
In Trichosurus and the Wombat, with immovable mandibular
rami, and in Phalanger, with the same only a little movable, the
angle of the mandible is broadly expanded on both sides for
the purpose of giving the powerful muscles a wide area of
insertion.
It has been stated above that the Koala and Pseudochirus, and
in a somewhat smaller degree Zrichosurus and Phalanger, must
move their mandibles in a lateral direction in the act of chewing,
so that the mandibular molars come quite opposite those of the
maxillary. This movement must take place in such a manner
that the whole mandible makes a slight lateral rotation with the
condyle of the same side, towards which the movement is directed,
as a fixed point or pivot. This rotation is effected by the con-
traction of a portion of the masseter of the opposite side, that
portion which, posteriorly, is inserted to the outer angle of the
mandible, and anteriorly to the foremost part of the zygomatic
arch or to the maxillary below and in front of the same.
When contracting, this muscle endeavours to pull its posterior
oint of insertion forwards, which results in a pressing of the
whole mandible towards the other side. The result is the more
easily obtained the more the outer angle of the mandible is
developed, because the lever becomes longer when the posterior
point of insertion of the said masseteric portion is situated more
laterally—or, which is the same, at a greater distance from the
pivot (condyle), The m. pterygoideus internus when contracting
endeavours to draw up the lower margin of the mandible, or its
inner angle, in an upward and somewhat median direction. This
action then results in an outward bending of the molar series,
and, perhaps, also a slight lateral rotation of the whole mandible,
because it is fixed posteriorly. The more the inner mandibular
angle protrudes in a median direction, the longer is the lever for
1902.] ADAPTATIONS IN DIPROTODONT MARSUPIALS. 31
this muscular action. It is thus possible that the movement of
the mandible towards one side is effected by the combined action
of a part of the masseter of the opposite side, and in less degree
by the pterygoideus internus of the same side. When the man-
dible has come to the desired lateral position so that the outer
margin of its molars comes quite opposite that of the corresponding
molars of the maxillary, both masseters and pterygoidet contract
and press the jaws together. The lower jaw then glides back
into the normal position, owing to the direction of the sloping
surfaces of the molar crowns as described above.
In the true Phalangerids the mandibular condyle which serves as
the pivot in the above-mentioned lateral rotation of the lower jaw
is steadied behind by the very solid post-glenoidal bone. In the
Wombat the condyle is only steadied on the inner side. We may
conclude from this that the chewing of this animal takes place
in a different manner from that in the true Phalangerids. The
sloping of the molar surfaces is also quite different in the Wombat,
i.e. inwards in the lower and outwards in the upper jaw. In
addition to this they are concave with sharp enamel ridges all
round, but especially protruding at the inner and outer margins,
It seems as if the chewing were effected by alternating trans-
verse movements of the lower jaw, and that the triturating action
on each side chiefly takes place when that side of the mandible is
moved in a median direction. As my material is not sufficient,
however, I shall not make any detailed statements.
Tarsipes takes, with regard to its dentition, the same extreme
position as it does with regard to the development of its intestine,
The mandibular incisors are slender and form together a point,
which perhaps might be used in making holes in the corolla of
flowers rich in honey for the insertion of the tongue. Together
with the upper incisors and canines they may also act as a pair
of pincers, useful when the animal catches small insects as it is
reported todo’, But the well-known rudimentary condition of
its molar series—in the specimen before me 2/2 to the left, 3/3
to the right—as well as the weakness of the lower jaw, w ithout a
processus coronoideus and angular inflection, make chewing or
even crushing of any hard prey impossible.
Thus the development and structure of the dentition, as well
as of the intestine, show a beautiful correspondence with the
diet and habits of the animals in the whole family Phalangeride,
the more striking through the polymorphism within so rece ere
limits as those of such a “natural group.
1 Conf. Lydekker’s ‘ Handbook,’ p. 121 (quoted on p. 18).
32 DR. L, V. LORENZ ON THE [Jan, 14,
3. On the Specimen of the Quagga in the Imperial Museum
of Natural History, Vienna. By Lupwie v. Lorenz,
C.M.Z.S.
[Received November 25, 1901. ]
(Text-figure 7.)
In the Zoological collection of our Museum there is a striped
Equus named “ Hquus quagga,” and until recently I have always
thought it was a Quagega of typical features, though the published
figures of that now extinct animal are rather different. But
when I visited the museums of Munich, Tring, London, Paris,
and Berlin last year, I discovered that the Quaggas which I saw
there were not quite in accordance with the specimen at Vienna.
I noticed them to be in general of somewhat different coloration—
more greyish or chocolate-brown on the upper parts, to have
narrower and perhaps more numerous dark stripes separated by
comparatively broader light interspaces, and, moreover, they: all
appeared to be of a smaller size. When I returned to Vienna I
asked my friend Marktanner (of the Museum in Graz) to photo-
graph our Quagga, and I had intended to send copies of the
photograph to different museums and to get others of the Quaggas
there in exchange. But different circumstances prevented me
from following the matter up until October last, when I had the
pleasure of receiving a visit from Dr. P. L. Sclater; and one of
his first questions was, what I thought about our Quagga, as it
seemed to him not quite identical with other specimens of this
Equus known to him. It was a great satisfaction to me that
such an authority as Dr. Sclater had come to the same conclusion
as I had done, and I am following his invitation in offering
to the Zoological Society of London an exact description of our
Quagga accompanied by one of the before-mentioned photographs.
Before writing this I examined the following figures of the Quagga,
which I propose to refer to as I proceed with my description :
Fig. I.—Buffon’s and Shaw’s copies of Edwards’s plate
(Gleanings of Nat. Hist. i. pls. 222 & 223)',
though this figure seems to me to represent
rather Hquus burchelli.
Fig. I1.—Buffon’s and Schreber’s copies of Allamand’s young
Quagga. (Allamand’s edition of Buffon, Suppleé-
ment, v. pl. vi.)
Fig. I1I.—Geoftroy St.-Hilaire and Cuvier’s plate (Hist. Nat.
Mammif. pl. 320), also reproduced by Schinz
(Siiugethiere, “‘ Hqwws,” pl. v.).
Fig. IV.—Schreber’s plate (vol. vi. pl. 3174), representing
the Quagga of Munich acquired by Ecklon about
1835,
1 Taken from the type of Equus quagga.
1902. ] QUAGGA OF THE VIENNA MUSEUM. 33
Fig. V.—The woodcut in Flower and Lydekker’s ‘ Animals
Living and Extinct,’ p. 384, fig. 160; copied in
W. L. Sclater’s ‘ Fauna of S. Africa,’ Mammals,
ii. p. 295, fig. 74.
Fig. VI.—A woodcut in Brehm’s ‘ Thierleben, Saugethiere,’
Bd. ui. fig. p. 49.
Fig. VII.—Noack’s drawing (Zool. Garten, 1893, fig. p. 293),
taken from the figures in the ‘ Gleanings of the
Knowsley Menagerie,’ pl. liv.
Fig. VIII.—The woodcut in Lydekker’s ‘ Royal Natural History ’
(vol. ii. fig. p. 507), perhaps representing the
specimen now in the British Museum.
Fig. IX.—The portrait of a Quagga’s head in Bryden’s ‘ Great
and Small Game of 8. Africa,’ pl. 11. fig. 2.
Fig. X.—Copies of York’s photographs (P. Z.8.1901, 1. p. 166,
fig. 47) of the Quagga in the Society’s Gardens
in 1870, and two photos of the same animal kindly
sent to me by Dr. Sclater.
Fig. XI,—An original photograph of the Quagga in the
Museum at Tring, being the same individual as
the last (no. x.).
Text-fig. 7.
The Quagga of the Vienna Museum. (From a photograph.)
Proc. Zoot. Soc.—1902, Vou. I. No. ITI. 3
34 DR. L. V. LORENZ ON THE
Description of the female Quagga
purchased by Ecklon, 1836 (Mus.
Vindob.).
Measurements in centimetres :
Total length from upper lip to
end of tail, without hair ... 300
Length of the face from
nostrils to the beginning of
PeWMANC Hance oenaaee 43
Length of the mane............ 76
From end of mane to root of
GAIN Saves Aa aan Manee sem A eeeaaas 128
anliwavhoutshangees se eeececee 40
{Bey \yatiln INEWOP scoodconoaneoosuus 80
SiG Ve vat aunt ties MH Nee DE aR 225
Ditto on the inner side ...... 15
Height at the withers......... 130
Height at the croup........ geben Lae)
Fore leg from the elbow ...... (al
Hind leg from the heel
loots jlenebheasn.cs es 6:5 & 6
Circumference of hoofs... 23 & 27
Coloration :
Ground - colour of upperside
clay-brown on the head, creamy
buff on the neck, shoulders, and
back, gradually changing to buff
on the flanks and thighs.
Breast, underparts, legs, and tail
white. Tail with elongated hair
from the root. Head, neck, back,
and flanks with narrow or broad
stripes of yellowish brown passing
into chestnut or maroon. Back
(haunches) clouded with drab.
Mane in the middle dark chestnut,
ornamented laterally by tufts of
whitish hair, ten on each side.
Along the back in continuation
of the mane a dark brown stripe,
having a breadth of 3 cm. on
the withers, expanding to 12 cm.
on the crupper, and growing again
narrower towards the tail,on which
it extends toa length of 12 cm.,
terminating with a breadth of
[Jan. 14,
_ Remarks on the Description.
Of all the Quaggas figured as
above noted the authors give gene-
rally smaller measurements, and
the specimens examined by me
were all apparently smaller.
Of the figures above cited only
figure iii. comes generally near the
colour of our specimen, but it is
still lighter. Fig. iv. approaches
it too, but is darker on the back.
In fig. i. the ground-colour is pale
chestnut.
The stuffed specimens seen in
other museums resemble in their
ground-colour fig. ii.
The stripes on the head, neck,
and body are darker except those
of fig. ili. Edwards's Quagga is
described as with black stripes.
In figure x. this band is only to
be seen on the croup; the pectoral
region of the spine appears to be
quite light® Figure 11. shows no
dorsal band. The young Quagga
of the Cape Town Museum is said
to be also without this band.
1902. |
0°5 cm. only. This dorsal band is
bordered on the pectoral region by
longitudinal creamy spots, which
become more confluent on the
lumbar and sacral parts, forming
at last continuous undulated lines
which vary in breadth from 0-5 to
15 cm.
A ventral band, beginning with
a brownish shade on the fore breast,
extends as a dark brown stripe to
the umbilicus: its greatest width
in the middle of the breast is about
8 cm., it narrows to 2 em. on the
belly.
Fetlocks with blackish rings just
above the hoofs. Sars creamy,
brownish at the back in the base,
dark clay-colour thence to the
end, extreme tips white.
Back of the nose nearly uniform
clay-colour, between the nostrils
dark brown ; lips whitish, chin and
throat uniform chestnut.
The dark striping is as follows :—
Hight narrow lines run from be-
tween the eyes down to the back
of the nose and up to the begin-
ning of the mane ; from the middle
of the front a ninth medial line
runs to the back of the nose.
From the eyebrows six stripes on
each side pass to the top of the
head. From the inner corner and
lower lid of each eye three in-
distinct stripes pass towards the
nose. On the cheeks upwards
from the corner of the mouth are
five streaks more or less curved.
Next to them four others on each
jaw, of which the first makes an
angle towards the eye, turning
then upwards to the base of ear ;
while the next two run more
directly in this direction, and the
fourth embraces the base of ear,
ending behind it at the mane.
The 2nd and 3rd of these stripes
are divided on the left side, the
6th on this side corresponding to
the 4th on the right side.
QUAGGA OF THE VIENNA MUSEUM. 35
The lateral spots or lines are
well marked in figures iv., vil., and
vii. only. Edwards’s figure (i.) has
white along the sides of the spinal
band with black spots on it.
This is said to be not present in
the young specimen at Cape Town.
These black rings are not to be
seen in figures 1., 11., 111., iv., and vi.,
besides being indistinct in others
(x.), but they appear on the photo-
graph (fig. xi.). White tips are not
observable on fig. ix. The back of
the nose is apparently dark in
figures iv., Vl., Vil., Vill.
The dark stripes of the head
appear very different in the various
figures. In fig. i. they are few in
number and very narrow, and the
interspaces are broad.
3*
36 ‘DR. L. V. LORENZ ON THE
On the neck there are eight
transverse bands, their respective
breadths being 2, 3-5, 4°5, 5°5, 5:5,
7, 5, and 3°5 on the right; 2°5, 3,
4-5, 6, 7:5, 9, 8, 4:5 on the left
side. The first six of them keep
more or lessapart, while the seventh
and eighth unite in front of the
fore-neck. The light creamy inter-
spaces on the neck are very narrow,
1 or 2 cm. only.
From the withers there run first
two stripes to the front of the
breast, where they join; they are
rather narrow above and grow
wider beneath. Then asingle stripe
that might be called a ‘“ shoulder-
stripe” also takes its origin from the
withers, and, passing the shoulders,
divides into two branches on the
humeral region. Inside the angle
thus formed are some irregular
and less distinct short stripes,
of which four or five are directed
obliquely upwards and _ partly
unite with three others directed
downwards and backwards. On
the body there are seven other
distinct bands getting more obso-
lete at the lower bifurcated ends,
and confluent at last with the buff
ground-colour of the flanks. Of
these the first three connected
with the longitudinal dorsal band
havea breadth of from 6°5 to 8 and
from 9 to 10 cm., the interspaces
between them being | and 1°5 em.
The fourth of these bands sends
an oblique branch to the croup,
and thus encloses a triangular
area of which the dorsal stripe
forms aside. Within this there
is another broad longitudinal stripe
anastomosing twice or three times
with the oblique one and also with
the dorsal stripe.
The triangles on both sides form
[Jan. 14,
These eight bands and the fol-
lowing two, or the homologues of
them, are to be recognized in most
of the figures, but they are in
general narrower and the inter-
spaces are broader. Fig. 111. comes
in this respect nearest to our
Quagga, and also does the colour.
The stripes on the head and neck
in figs. iv. and 1x. are much darker.
Edwards’s Quagga (fig. i.) shows
unusually narrow black bands on
the neck and broad interspaces, just
the contrary to our Quagga.
The bifurcation of the shoulder-
stripe is well seen in most of the
figures except in fig. iv.
These stripes are not repre-
sented in most of the figures.
The vertical body-stripes are
different in every figure. On the
whole they are narrower and more
numerous, besides they do not ex-
tend to the haunches. i
These oblique stripes are not to
be seen in some of the figures (i11.,
iv., and x.), in others there are
spots in their places (figs. i. and
Vil.)
The saddle is wanting in figs. i.
1902.]
together a kind of saddle, as is the
case in all striped horses of the
burchelli-group. The fifth band
takes an oblique direction through-
out, running as well as the sixth
over the haunches, both becoming
gradually narrower at their upper
ends, and not quite reaching the
dorsal band.
A seventh, somewhat narrower
but still distinct, although twice
interrupted, stripe takes a direc-
tion from the groin and goes over
the haunches to the root of the tail
without reaching it. Between the
6th and 7th stripe is an indistinct
short band. Three or four other
obliqueand gradually fading stripes
are observable on the back of the
haunches.
QUAGGA OF THE VIENNA MUSEUM. oe
to iv. It may be recognized on v.
and vi. and on the photo (xi.), as
also the bands on the quarters,
but they are not seen in fig. x.,
which represents the same indi-
vidual.
This reminds one of Hquus
burchelli.
On comparing the stripes and bands of our Quagga with the
pictures of the other Quaggas and with the various forms of the
Zebras of the Burchell-group, there seems to me no question
that the Quaggas belong to that group. Jalso have the impression
that, in spite of the variability of the marking, the examination
of sufficient material would result in ascertaining the existence
of homologous stripes in the group above mentioned.
From a
further careful comparison of all the different figures, and
especially of the original picture of Edwards, with the stuffed
specimens, or at least with photos of them, we could perhaps
obtain sufficient answers to the following questions :—
(1) Is the Vienna Quagga specifically the same as Edwards’s
Quagga ?
(2) Can other so-called Quaggas (as, for instance, those of the
British Museum and of the Tring Museum) be identified with
Edwards’s Quagga, notwithstanding the differences pointed out
so exactly by Mr. Pocock? (Ann. Mag. Nat. Hist. ser. 6, xx.
p. 37).
(3) Can the Vienna Quagga be identified with the Quaggas of
London and Tring ?
To these questions I would only reply provisionally that the
differences between Edwards’s picture and the Vienna, London,
Tring, and other specimens are certainly more essential than the
differences between the Vienna Quagga on one side and the
London, Tring, and other Quaggas on the other.
Kdwards’s
Quagga, as already remarked, much resembles Hquus burchelli in
some respects—e. g., in the black stripes, well defined on the head
and extremely narrow on the neck, and in the tufted tail.
As to the Vienna specimen, it is possible that its characters
38 MR. J. L, BONHOTE ON [Jan. 14,
may be merely individual, for we find among skins of Zebras from
the same locality some with pure black stripes and others with
brownish stripes. Besides, the transverse stripes on the body of
our Quagga show a tendency to bipartition, and the oblique
stripes incline to break up into blotches. There likewise remains
the possibility that our specimen has been rather increased in
size by the art of the taxidermist. Considering, however, that so
many local forms of Equus burchelli have been distinguished
during the last few years, it is by no means impossible that the
Vienna specimen might be ultimately separated subspecifically
from other Quaggas.
Vienna, Nov. 20th, 1901.
4, On a further Collection of Mammals made by Mr. Th. H.
Lyle in Siam. By J. Lewis Bonnors, M.A.
[Received November 19, 1901.]
The following paper gives an account of a further small consign-
ment of Mammals sent home by Mr. Th. H. Lyle from Siam.
Although small in point of numbers it contains several specimens
of considerable interest, and foremost among these is a fine
example of the Siamese Hare, which proves to belong to a
species not hitherto described. A specimen of Sctwrus atro-
dorsalis, in immature pelage, and two specimens of Mustela
flavigula form a valuable addition to the National Collection, and
help considerably to the more correct understanding of their
respective groups.
1. Cynoprervus spHinx (Vahl).
Vespertilio sphinw Vahl, Scrivter af Naturhistorie-Selskabet,
Ate Band, 1%* Heft, p. 123 (1797); Bonh. P. Z.S. 1900, p: 191;
id. loc. cit. p. 875.
Cynopterus marginatus (Geoftr.), Flower, P. Z.S. 1900, p. 341.
a. 2. N. Chiengmai, 27th Feb., 1901.
9. MUSTELA FLAVIGULA Bodd.
Mustela flavigula Bodd. Elench. Anim. p. 88 (ex Penn.) (1785) ;
Flower, P. Z.8. 1900, p. 333.
Mustela flavigula subsp. typica Bonh, Ann. & Mag. Nat. Hist.
ser. 7, vol. vii. p. 344 (April 1901).
a,b. &. N. Chiengmai, 28th Feb., 1901.
These two individuals closely agree with the description in my
paper quoted above, with the exception that the hind-quarters
could hardly be styled “very dark brown”; this apparent dis-
crepancy is, however, merely due to faded pelage, for of the two
specimens one is lighter than the other.
IT append the measurements taken in the flesh, as they are
~
1902.] MAMMALS FROM SIAM. 39
slightly at variance with those taken from the dried skin in my
former paper :—
Head and Tail Hind foot Fa
body. without hairs. (s. u.). ue
Gea son 589 mm. 440 mm, 108 mm, 4 mm,
fee 565 ,, 435 ,, Oe ew
3. SCIURUS CASTANEOVENTRIS GORDONI Anders.
Sciurus gordont Anders. P. Z.S. 1871, p. 140; id. Zool. Res.
Yunnan, p. 240 (1879).
Sciurus castaneoventris gordont Anders., Bonh. Ann. & Mag.
Nat. Hist. ser. 7, vol. vii. p. 164 (Feb. 1901).
a. 3. Doi Sritepe, Chiengmai, 27th March, 1901.
Dimensions in flesh. Head and body 218 mm.; tail 193; hind
foot 47; ear 21.
This form has hitherto been recorded only from Upper Burma,
but the present specimen agrees perfectly with examples from the
type locality.
4, ScruRusS CANICEPS Gray.
Sciurus caniceps Gray (nec Temm.), Ann. & Mag. Nat. Hist.
x. 1842, p.263; Bonh. P.Z.S. 1901, p.55; id. Ann. & Mag. Nat.
Hist. ser. 7, vol. vii. p. 271 (March 1901).
a,b,c. 629. Sawankalok, Siam, 27th Dec., 1900.
Two of these specimens are passing into the bright pelage, while
the third has fully assumed it.
5, ScruRUS ATRODORSALIS Gray.
Sciurus atrodorsalis Gray, Ann. & Mag. Nat. Hist. x. 1842,
p. 263; Bonh. P.Z.8. 1901, p. 55.
a @imm. Chiengmai, Siam, 5th April, 1901.
This specimen, which is about three-fourths grown, differs from
the adult in its much greyer coloration, the annulations on each
hair being of a very pale grey. The colouring of the ears, however,
shows a faint yellowish tinge, and down the centre of the back
there isa slight trace of the dark colour characteristic of the adult.
The underparts resemble those of the adult.
6. Mus concotor Blyth.
Mus concolor Blyth, J. A.S. B. xxviii. p. 295 (1859); Bonh.
P. Z. 8. 1900, p. 195; Flower, loc. cit. p. 361.
a. gad. Chiengmai, Siam, 3rd April, 1901.
b. Qad. Chiengmai, Siam, 21st April, 1901.
7. Mus serpont (Blyth).
Leggada jerdont Blyth, J. A.S. B. xxxii. p. 350 (1863).
a. 6 ad. Doi Sritepe, Chiengmai, 15th April, 1901.
b,c. @ ad. Doi Sritepe, Chiengmai, 16th & 17th April, 1901.
40 ON MAMMALS FROM SIAM. [Jan. 14,
8. LEPUS SIAMENSIS, Sp. n.
Lepus sp. inc. Flower, P. Z.8. 1900, p. 365; Bonh. P.Z.8.
1901, p. 56.
General colour above fulvous and dark brown, the latter colour
beconiing absent on the hind-quarters and flanks, where the
fulvous is slightly tinged with rufous. The whole of the under-
parts except the lower neck and chest pure white, the line of
demarcation being sharply defined. The neck, chest, and limbs
are fulvous of varying shades, the colour being deepest on the
fore legs, where it is tinged with rufous, and palest on the inner
sides of the hind limbs, where it becomes nearly white. Hach
hair is dull white or greyish at-the base, shading into dark brown
(seal-brown, Ridgw.') and having a broad subterminal fulvous
(buff, Ridgw.) annulation.
On the head the fulvous becomes deeper in colour, and there is
an ill-defined whitish stripe running from the nostril to the front
of the eye on either side.
The ears, which are but scantily clothed with hair on their
outer posterior surface, resemble on the anterior surface the
general colour of the back, although the darker tint predominates.
The outer anterior and posterior margins are white. At the tip
the inner surface is clothed with pure fulvous hairs, while on the
external surface the hairs are dark brown.
The tail is dark brown above throughout its length and white
underneath, with a slight tinge of buff on the sides.
The skull resembles most nearly that of Z. peguensis, from
which it differs chiefly in the muzzle being slightly broader at its
base. The basioccipital bulges outwards and downwards on
either side instead of having its sides parallel, thus causing the
bulle to appear at first sight somewhat smaller. The skull as a
whole is, moreover, rather larger. Comparing the grooves on the
upper incisors with those figured in Dr. Forsyth Major’s paper
(Trans. Linn. Soc., 2nd ser. Zool. vol. vii. p. 468, 1899), it appears
to be most nearly allied to Z. hainanus, although somewhat in-
termediate between it and Z. dayanus. The groove in the species
under consideration is moderately broad and nearly rectangular,
with a small rounded process jutting out at about the centre of
the outer margin.
Dimensions of type (in flesh). Head and body 435 mm.; tail
66; hind foot 95; ear 82.
Skull. Greatest length 86; breadth of palate at Ist molar 13;
length Ist premolar to outer edge of incisors 27; greatest breadth
of brain-case 30.
Hab. Siam.
Type. B.M.1.7.7.13. 3 ad. Chiengmai, 16th Feb., 1901.
This fine species is most nearly allied to LZ. haimanus Swinhoe,
from which it is easily distinguished by its greater size and the
1 Ridgway, ‘ Nomenclature of Colours,’ Boston, 1886.
1902. | ON RHYNCHOTA FROM UGANDA. 4)
absence of a clearly defined white supra-orbital stripe. From
LL. peguensis, to which it more nearly approaches in size, it differs
in the fur on the back and tail being dark brown instead of
black, and in the absence both of the ashy tinge on the rump
and the black terminal patch on the posterior outer surface of
each ear. ;
The skull of a species of Lepus sent home by Mr. Lyle in a
former collection agrees with the type skull. The animal to
which it belonged was unfortunately destroyed, but Mr. Lyle
writes that it was a female, and the following are the measure-
ments in the flesh :—Head and body 463 mm.; tail 74; hind foot
97; ear 84,
For the sake of comparison the measurements of the skulls of
L. hainanus (type), L. peguensis, and the two skulls of this species
are appended :—
et, | Tepws | Tepus | Zepus
(type). | Stamensis.| hainanus. | peguensis.
mm. mm mm. mm.
Greatest length ...........:0cccc000s secs 86 89 72 85
Breadth between 1st molars ............ 13 13 il 13
Least breadth between orbits ......... 13 12 12 13
Length from 1st premolar to outer 26 27 21 25
edge of incisors.
Greatest breadth of brain-case ......... 30 28 26 28
Height, crown to base of lower jaw ... 54 ee He 53
Greatest breadth of basioccipital ...... 10 11 10 9
Posterior breadth of nasals ............ 21 ce 16 18
5. On the Insects of the Order Rhynchota collected by
Sir Harry Johnston, K.C.B., in the Uganda Protec-
torate. By W. L. Disrant.
[Received November 23, 1901. |
(Text-figure 8.)
The few specimens of this Order collected by Sir H. H. Johnston,
and by him presented to the British Museum, are principally
interesting as showing that the Uganda Rhynchotal fauna and
that of West Africa are practically identical. The species known
only from East Africa are very few, and further knowledge may
prove them still fewer. Two new species are described, one of
which has a far wider distribution than the Uganda Protectorate.
I have added notes to the enumeration of each species as expla-
natory of its geographical dispersion,
AQ MR. W. L. DISTANT ON [Jan, 14,
HETEROPTERA.
Fam. PENTATOMIDA.
Subfam. ScUTELLERINA.
CRYPTACRUS COMES.
Tetyra comes Fabr. Syst. Rhyng. p. 130. 8 (1803).
Var. Dist. Ent. Monthl. Mag. xiv. p. 75 (1877).
Mt. Ruwenzori.
Resembling the variety I described from the Camaroons (supra),
but with slight traces of an ochraceous subapical fascia to the
scutellum. A highly variable and widely distributed species
found all over tropical and subtropical Africa.
Subfam. DrnrpDorin&.
CYCLOPELTA TRISTIS.
Dinidor tristis Stél Hem. Afr. i, p. 212 (1864).
Mts. Ruwenzori and Entebbe.
A species hitherto known only from West Africa.
ASPONGOPUS LIVIDUS.
Aspongopus lividus Dist. Ann. Mag. Nat. Hist. (7) xi. p. 315
(1898).
Var. Abdomen above testaceous.
Mt. Ruwenzori.
In typical specimens described from Nyasaland the abdomen
above is dark olivaceous. I can, however, discover no other differ-
ential characters.
ASPONGOPUS NIGRO-VIOLACEUS.
Pentatoma nigro-violacea Pal. Beauv. Ins, p. 83, Hem. pl. 7.
fig. 4 (1805); Dist. Ent. Month. Mag. xv. p. 10 (1878).
Mt. Ruwenzori.
A species hitherto recorded only from West Africa.
Fam. COREID A,
Subfam. Micrin az.
ANOPLOCNEMIS TRISTATOR.
Lygeus tristator Fabr. Syst. Rhyng. p. 206 (1803).
Mt. Ruwenzori.
A West-African species.
Fam. PYRRHOCORIDA.
Subfam. PyrrHocoRIn@&.
ODONTOPUS NOTABILIS, sp. n.
Ochraceous ; antenne black; head, basal joint and base of
1902. ] RHYNCHOTA FROM UGANDA. 43
second joint of antennee, and legs reddish ochraceous ; scutellum,
base of clavus, a round spot near apex of corium, discal area of
prosternum, anterior areas of meso- and metasterna, a lateral
basal spot behind eyes, two discal transverse lines to pronotum
(one curved near anterior margin the other straight near centre),
and abdominal segmental incisures (not extending to lateral margins
and concolorous on disk), black.
Text-fig. 8.
Odontopus notabilis.
First joint of antenne with a few distinct hairs near base and
some hairs at apices of second and third joints, third joint
shortest, second and fourth joints subequal in length; corium
and clavus very thickly, finely, and obscurely punctate; the
black base of clavus coarsely punctate.
Long. 16 to 22 mm.
Entebbe.—The British Museum also possesses specimens from
Kast Central Africa (G@. F. Scott Elliot); Kavala Island, Lake
Tanganyika (4. Carson), and Angola.
SERICOCORIS JOHNSTONI, Sp. n.
Head, pronotum, scutellum, body beneath, and legs ochraceous :
lateral margins of pronotum, the corium, and lateral margins of
sternum pale purplish antennz, central longitudinal fascia and
basal margin to head, the margins of the anterior area of pro-
notum, basal margin of scutellum, rostrum, tibie, tarsi, and
margins of sternal segments, black; membrane pale brownish.
Pronotal margins somewhat strongly reflexed ; corium and clavus
thickly punctate.
Long. 14 mm.
Entebbe.
a4 DR. A. G, BUTLER ON [Jan. 14,
Fam. REDUVIID.
Subfam. ACANTHASPINS,
PLATYMERIS RHADAMANTHUS.
Platymeris rhadamanthus Gerst. in y. d. Deckens’s Reise, iii.
(2) p. 419, pl. xvii. fig. 8 (1873).
Baringo, 4000 ft.
A species known only from East Africa.
HOMOPTERA.
Fam. C1IcADIDS. ,
PLATYPLEURA RUTHERFORDI. ue»
Platypleura rutherfordi Dist. Ann. Mag. Nat. Hist (5) xi.
p. 173, pl. ii. fig. D (1883).
Entebbe.
A species originally described from West Africa, and since
received from Mashonaland.
Fam. CERCOPIDS.
Subfam. APHROPHORIN.
PTYELUS FLAVESCENS. 2
Tettigonia jlavescens Fabr. Ent. Syst. iv. p. 24. 30 (1794).
Entebbe.
A most variable species distributed over the greater part of
tropical and subtropical Africa.
6. On two Collections of Lepidoptera made by Sir Harry
Johnston, K.C.B., in the Uganda Protectorate during
the year 1900. By Arruur G. Burtsr, Ph.D., F.LS.,
F.ZS., &. ; Senior Assistant-Keeper, Zoological De-
partment, British Museum (Nat. Hist.).
[Received November 12, 1901.]
(Plate I.*)
The two collections of which the following is a list were obtained
at Entebbe, Port Alice, Port Ugowe, Busiro; and from Ruwenzori,
Toro, and the Congo forest, respectively. The first consignment
consisted chiefly of well-known and widely distributed forms ;
but the second not only included a good sprinkling of rarities, of
species new to the Museum Collection, and even of undescribed
species, but was especially interesting from the strange com-
mingling of Eastern and Western types which it contained.
1 For explanation of the Plate, see p. 51.
1h is) ALCO poll I, teak IL,
Horacelimght delet ith West Newman chromo.
LEP LOOP TERRA FROM WGAN DA.
1902.] LEPIDOPTERA FROM UGANDA. 45
Among the rare species, Melinda mercedonia, Monotrichtis
saussuret, Ypthima albida, Charaxes bipunctatus, Harma hobarti,
Diestogyna amaranta, Neptis wicomedes, Acreea toruna, Acrea
orinata, Acrea oreas, Terias punctinotata $, Belenois solilucis,
Belenois raffrayi, Papilio lormieri, and Celenorrhinus opalinus 2
are worth special note. The new species are Harma johnston,
Pseudathyma plutonica, and Aphneus hollandi: there is also a
new moth.
The following is a list of the species :—
1. Amauris niavius Linn. Toro, June 16th.
2. is enceladus Brown. 5 ns
3. % albimaculatus Butl. _,, Ms
4. Melinda mercedonia Karsch. ,, -
5. Tirwmala petiverana Doubl. .
6. Monotrichtis perspicua Trim. Ruwenzori, 5000 ft., Sept.
(le 44 safitza var. campina Auriv. Ruwenzori ,4200 ft.,
Sept.
8. saussuret Dewitz. Ruwenzori, 000 ft., Sept.
ws 2G pele 6 granulosa Butl. me bs aS
10. Fs albida Butl. -
11. Charaxes numenes Hewits. Entebbe, April 30th.
os tiridates Fabr. Toro, June.
13. BS bipunctatus Roths. ,,
14. Precis boopis Trim. Port Ugowe, 20th Feb. & 23rd J uly.
la, —y, «| clea Cram. . 20th & 21st Feb.
16. ,, cebrene Trim. oe 20th & 22nd Feb.
7. 4, westermanni Westw. Toro, June.
» terea Drury. Entebbe, April 20th.
19. 4, -yregorw Butl. Toro, June 16th.
20. chorimene Guér. Port Ugowe, April 21st & 22nd.
2 Pr. oto goniomorpha temora Feld. Tor o, June 16th.
22. Hypolimnas salmacis var. Drury. Congo forest, July 16th.
23. a3 misippus Linn. Port Alice, 23rd March.
24. Chloropea lucretia Cram. Toro, June.
25. Harma johnstoni Butl. Toro, June 16th.
26. - cents Drury. Congo forest, July 4th.
27. + ,, hobarti Butl. Toro, June 16th.
28. ,, aramis Hewits. Congo Forest, July 16th.
29. theobene Doubl. = ~
30. Orenis occidentalium Mab. Busiro, June2nd; Toro, June 16th.
31. ,, boisduvalia Wilgr. as M
32. Huphedra eleus vav., Drury. Congo forest, J uly.
30. M imanum vain. ., Butl. Toro, June; Congo forest,
: July 16th.
34. 7 cypetina Stgr. Congo forest, July 16th.
35. Bs spatiosa var., Mab. ‘Toro, June.
36. Aterica galene Brown. Congo forest, July 16th.
37. Cynandra opis Drury.
38. Huryphene abesa Hewits.
7 ”
”? 99
DR. A. G. BUTLER ON [Jan. 14,
. Diestogyna amaranta Karsch. 2, Toro, June 16th; ¢, Congo
forest, July 16th.
p. 2 (9 near felicia). Congo forest, July 16th.
E Bywdatiioens plutonica Butl. Toro, June.
. Kallima rumia Westw. Toro, June 16th.
. Hurytela hyarba Fabr. Busiro, 5050 ft., June 2nd.
. Ergolis enotrea Cram. Toro, June 16th.
. Catuna crithea Drury. _,, ns
. Neptis nicomedes Hewits. Busiro, 5500 ft., June 2nd.
» melicerta Drury. Congo forest, July 16th.
| Atella phalantha Drury. Port Alice, March 20th ; Entebbe,
April 30th.
. Acreaztoruna 2, Gr.-Sm. Toro, June 16th.
,, alicia Sharpe. %s 3
» uvui Gr.-Sm. = $
» vinidia Hewits. F ee
» serend, var. rougetia Guér. Port Ugowe, Feb. 20th.
- 20980000, var. lycia Fabr. Entebbe, April 20th.
,» onerata Trim. Port Ugowe, Feb. 20th & 22nd.
» ndatalica, var. dissociata Gr.-Sm. Ruwenzori, 7000 ft.,
Sept.
» setes, var. menippe Drury. Entebbe, March 20th &
April 30th.
» orinata Q Oberth. Entebbe, April 30th.
oreas Sharpe. Toro, June 16th.
; Megalopalpus 2 zymna Westw. Congo forest, July 16th.
. Zeltus? antifaunus Hewits.
. Aphneus hollandi 3 Butl.
. Cacyreus lingeus Cram. ‘Toro, June 16th.
. Azanus natalensis Trim.
. Zizera antanossa Mab. Port Ugowe, Feb. 22nd.
. Vychitona ies var. aleesta Cram. Entebbe, April 30th.
oP) oP
» eummaculata Auriv. Toro, June 16th.
; Colias electo, var. edusa Fabr. Toro, June.
. Terias brigitta, var. zoe Hopft. Port Ugowe, Feb. 21st & 22nd.
» boisduvaliana Mab. ¥s s
» punctinotata § Butl. Toro, June.
Catopsilia florella Fabr. Port Ugowe, Feb. 21st & 23rd.
. Belenois solilucis Butl. | Toro, June 16th.
eS calypso var. , Drury.
- instabilis Butl. ° Port Ugowe, Feb. 20th; Toro,
June 16th.
» formosa Butl. Toro, June 16th.
- severina var. infida Butl. Port Ugowe, Feb. 20th
to 23rd.
3 mesentina Cram. Port Ugowe, Feb. 20th to 23rd.
» vraffrayi Oberth. Toro, July 16th.
. Pinacopteryx liliana Gr.-Sm. Port Ugowe, Feb. 23rd.
. Leuceronia argia 2, var. idotea Boisd. Congo forest, July.
a pharis 3, var., Boisd. Toro, June.
1902. ] LEPIDOPTERA FROM UGANDA. 47
82. Hronia dilatata Butl. Port Ugowe, Feb. 23rd.
83. Papilio policenes Cram. Port Alice, March 20th.
84. », demodocus Esper. Entebbe, April 30th; Toro
June 16th.
85. » lormiert Dist. ‘Toro, June.
86. Hretis perpaupera Holl. Toro, June 16th.
87. Celenorrhinus opalinus 2 Butl. ,, y
88. Baoris inconspicua Bertol. Entebbe, April 30th.
New Species, ce.
HARMA JOHNSTONI, sp. n. (Plate I. figs. 4, 5.)
Nearly related to H. herminia of Grose-Smith ; larger, with
almost the same pattern: the male paler and more olivaceous at
base; the pale yellow belt of the secondaries much wider; the
blackish macular belt across the disk of the wings rather wider
on the primaries and with its inner edge on the secondaries acutely
zigzag; the external area less ochreous; the irregular black
submarginal line better defined and with long denticles pointing
outwards on the folds between the nervures; the yellow lunate
band between this line and the discal belt considerably narrowed
and partly obscured by dark brown; the external border dark
brown, only interrupted by pale yellow patches on the subcostal
interspaces in the secondaries, but in the primaries interrupted
by small patches excepting on the lower radial interspace: on the
under surface the pattern is similar to that of H. herminia, but
the colouring is less rosy, greyer, the enclosed irregular band
limited externally by the straight central line is narrower and
becomes uniform with the ground-colour below the subcostal vein.
Expanse of wings 72 mm.
The female has the general pattern above of what I regard as
H. herminia Q (an insect nearly related to H. capella 92); the
primaries are, however, much more produced at apex and the
secondaries at anal angle; the basal area of the wings is much
more broadly suffused with ferruginous along the veins, the central
blackish band on the secondaries is almost obliterated, only clearly
discernible towards the costa, but is followed by a series of
indistinctly whitish-edged grey lunules followed by white dots,
the submarginal irregular black line being indistinct excepting
for the denticles on the folds between the veins: on the under
surface, as in the supposed female of H. herminia, the pattern
and general colouring nearly approach those of H. lurida 2 , but
with less white on the primaries and with the central line much
narrower and red-brown rather than brick-red. Expanse of
wings 89 mm.
Toro, 16th June, 1900.
DIEsTOGYNA AMARANTA Karsch. (Plate I. figs. 2, 3.)
9. Dark olive-brown above, irrorated with pale ochreous and
banded with the same colour; discoidal cell of primaries with
48 DR. A. G. BUTLER ON [Jan. 14,
similar pale-bordered but indistinct markings; a discal increasing
oblique belt slightly curved, with sinuated inner edge and diffused
outer edge from subcostal to first median vein, from first median
vein to inner margin abruptly narrowed and of equal width,
slanting obliquely inwards; a chain-like double series of opposed
lunules parallel to outer margin; outer border rather paler
(because more densely irrorated with pale ochreous) than the rest
of the wing: secondaries crossed from just before middle by four
pale ochreous bands, the first slightly irregular and sharply defined
internally, diffused externally, continuing the discal belt of the
primaries; the second and third continuing the chain-like series
of the primaries, the fourth submarginal, less defined than the
others, undulated ; outer border as in primaries. Wings below
altogether paler than in the male, greyer, the discal belt of upper
surface well defined, but pinky whitish on the secondaries ; the
basal area of these wings irrorated with pearl-grey indicating two
vague subbasal bands; the chain-like belt pearl-grey on both
wings and with white points on the upper internal lunules of the
primaries and the lower internal lunules of the secondaries.
Expanse of wings 56 mm.
Toro, June 16th.
Prof, Aurivillius has pointed out that the female figured
by Karsch does not belong to this species and has named it
D. karscha.
A second female similarly coloured is in the collection; but,
without the male, it would be rash to name it: in pattern it is
not unlike D. felicia, but it is a much shorter-winged insect.
PSEUDATHYMA PLUTONICA, sp. n. (Plate I. fig. 6.)
Allied to P. sibyllina, but smaller, shorter in wing, the primaries
with much less sinuous outer margin, the secondaries rounded,
not produced at anal angle; the discal belt of the primaries
forming three patches, the first three divisions being much shorter
than in P. sibyllina; the belt of the secondaries constricted
towards costa and not deeply indented externally; the inner
submarginal line only white in the centre: on the under surface
the markings on the basal area, excepting a costal patch on the
secondaries, suffused. Expanse of wings 42 mm.
Toro, in June.
ACR#A ORINATA Oberthiir. (Plate I. fig. 1.)
@. This is the largest female of the group yet described, and
much more nearly resembles the male of A. parrhasia than any-
thing else: the primaries show a pinky-white, semitransparent,
oblique, trifid bar beyond the cell in continuation of the discal
tawny belt, and the basal area of the secondaries is almost wholly
black: on the under surface this sex chiefly differs from A.
orinata S$ in showing a diffused whitish patch beyond the cell of
primaries. Expanse of wings 70 mm.
Entebbe, April 30th.
1902. } LEPIDOPTERA FROM UGANDA. 49
I once supposed that A. orinata would prove to be a variety
of A. oppidia $, and Prof. Aurivillius believed it to be a form of
A, orina, but we were both wrong; it is a good distinct species.
APHNEUS HOLLANDI, sp. n. (Plate I. fig. 7.)
3. Nearly allied to A. orcas; but the metallic colouring of the
upper surface more brilliant and rather emerald-green than
greenish blue; the black cell-spots on the primaries are consider-
ably larger and the apical area is black with scarcely a trace of
metallic scaling, the subapical series of spots (of which only the
two uppermost are clearly visible) reduced to a few metallic
scales; on the secondaries the metallic patch extends closer to the
outer margin, the apical area is browner, and the marginal spot
between the tails is ochreous instead of red: on the under surface
the differences are much more marked; the ground-colour of the
primaries is of a palish earthy brown with the silver markings
bordered with deep maroon; the arrangement similar to that in
A. orcas, but the short band at end of cell truncated in front and
gradually narrowing backwards; no submarginal silver spots ;
the oblique streak towards external angle very narrow; secondaries
with the ground-colour yellowish stone-colour suffused with grey
(or sordid) towards base and apex; the silver markings bordered
with ferruginous red; the arrangement of these markings is
similar to that in A. orcas, but the submarginal series is placed
upon a ferruginous band and is almost obliterated excepting at
anal angle; the oblique internal bar above the latter is curved, so
as almost to join the broad discal belt, and the two silver spots
above it are greatly reduced in size; the anal lobe is much paler
in colouring—ochreous with a quadrate central ferruginous patch ;
the fringe brown where it is black in 4.orcas. Expanse of wings
39 mm. i
Congo forest, July 16th, 1900.
I have named this beautiful little butterfly in honour of my
friend Dr. W. J. Holland of Pittsburg, whose admirable photo-
graphic plate in the ‘Entomological News’ for 1893 has greatly
facilitated the identification of the species of Aphneus.
BELENOIS CALYPSO Drury.
Var. 6. The secondaries white below, with the usual markings,
but the orange streaks at base and apex of costa (which are usually
ill-defined) and a dash at the base of the submedian vein sharply
defined in deep orange (more so than in B. dentigera).
Prof. Aurivillius correctly states that B. agylla is synonymous
with B. solilucis (not with B.ianthe). Until I saw the specimens
in the present collection, I was not aware that the border of
the primaries was ever so wide in BS, solilucis as is shown in
' Rogenhofer’s figure, and I naturally supposed the regularity of
the border in that figure to be due to inaccurate drawing.
» Proc, Zoou. Soc,—1902, Vou. I. No, IV. 4
50 ON LEPIDOPTERA FROM UGANDA. [Jan, 14,
BELENOIS FORMOSA Butl.
The intermediate phase of this species from Toro differs from
the wet phase in the smaller and partly obliterated white spots in
the apical border of the primaries; and on the under surface
in the pale earthy-brown subapical streak and veins on the primaries
and the brown veins and markings on the secondaries. The dry
phase (which we obtained from the Crowley collection) is still less
spotted on the apical area of the primaries above, has the apical
area of these wings below and the secondaries of a whity-brownish
tint with still paler brown markings than the intermediate
phase.
3, Toro (Sir H. Johnston); 3 3, Mt. Elgon (7. J. Jackson).
Among the Lepidoptera Heterocera there was nothing of
interest with the exception of a very remarkable new genus
of Sesiide, which Sir George Hampson has asked me to describe.
CRYPTOMIMA, gen. nov.
Allied to Ceratocorema. Wings for the greater part opaque,
brilliantly metallic: primaries narrow, elongated, the costa nearly
straight to 3, then gradually deflexed to apex, which is moderately
acute; outer margin very oblique, slightly convex, passing
gradually into inner margin, which is slightly concave almost to
the base: secondaries with the costal margin nearly straight
to apex, the apex moderately defined, the outer margin slightly
arched to first median branch and thence nearly straight to anal
angle; abdominal margin sinuous, widest in the centre. The
neuration may be characterized by veins 7 & 8 of the primaries
being emitted from a long footstalk; vein 4 of the secondaries
absent. Body smooth and shining; the antenne simple; palpi
rather slender, elongated, second joint upturned, slightly curved,
third joint porrected at an oblique angle, spine-like. Front legs
with tibie coarsely fringed below; second pair fringed externally,
the upper end of the joints with the fringe projecting, two well-
developed unequal spurs ; third pair smooth, with two long unequal
spurs beyond the middle and two below the end of the tibize, the
latter joints and the tarsi coarsely setose: abdomen with a long
densely scaled process with naked extremity from the dorsal surface
of the terminal joint, resembling the ovipositor in certain /ehneu-
monide ; vulva tufted.
CRYPTOMIMA HAMPSONI, sp. n. (Plate I. fig. 8.)
Wings above steel-blue glossed with green, brilliantly metallic :
primaries with a small bifid subbasal patch divided by the median
vein; end of cell and median vein blackish blue; beyond the cell
a hyaline belt brilliantly shot with golden-green from near costa
to near external angle, slightly increasing in width from front to
back of wing: secondaries with the basal two-sevenths hyaline
crossed by steel-glossed black veins. Body above steel-black,
1902. | THE SECRETARY ON ADDITIONS TO THH MENAGERIE, 51
slightly purplish; eyes red-brown; face opaline whity brown;
palpi whity brown, second joint with opaline white scales ; pectus
and legs steel-blue; terminal joints of hind tarsi whitish below ;
venter nacreous, the base broadly whitish followed by a still
broader dull steel-blue band, beyond this a second whitish band
or patch and then a second steel-blue band ; anal fringes ochraceous
and smoky black. Expanse of wings 35 mm.
Toro, June 16th, 1900.
I have named this remarkable insect in honour of my colleague,
Sir George Hampson, Bart., who is engaged upon a complete
Catalogue and Revision of the African Lepidoptera Heterocera.
EXPLANATION OF PLATE I.
Fig. 1. Acrea orinata &, p. 48.
. Diestogyna amaranta 3, p. 47.
- 47,
29 22 2 3
. Harma johnstoni 8, p. 47.
” ” » PD. 47.
. Pseudathyma plutonica 6, p. 48.
. Aphneus hollandi 6, p. 49.
. Cryptomima hampsoni 2, p. 50.
COT > OV GO LO
February 4, 1902.
Prof, G. B. Howss, LL.D., F.R.S., Vice-President,
in the Chair.
The Secretary read the following report on the additions to the
Society’s Menagerie during the month of January 1902 :—
The registered additions to the Society’s Menagerie during the
month of January were 87 in number. Of these 32 were acquired
by presentation and 3 by purchase, 47 were received on deposit,
4 were born in the Menagerie, and 1 was received in exchange.
The total number of departures during the same period, by death
and removals, was 168.
Amongst the additions attention may be specially directed to :—
1. A female White-tailed Gnu (Connochetes gw), born in the
Menagerie on January 10th, from one of the females presented
by Mr. C. D. Rudd, F.Z.S., in August 1901.
2. Nine Pheasant-tailed Jacanas (Hydrophasianus chirurgus)
from India, presented-by My. Frank Finn, F.Z.S8., on January 11th.
This peculiar bird is new to our collection, and we are greatly
obliged to Mr. Finn for sending us the specimens, as also to
My. Knifton, of the P. & O. s.s. ‘ Malta,’ under whose care they
were placed during the voyage home.
3. Three Red River-Hogs (Potamocherus penicillatus), born in
the Menagerie on January 27th.
The breeding of the Red River-Hog in captivity is a noticeable
4x
52 MR. F, E, BEDDARD ON A GIRAFFE. [Feb. 4,
event, but it has already occurred on two previous occasions in
the Society’s Menagerie (cf. P. Z.S. 1861, p. 62, pl. xii.).
Mr. F. E. Beddard, F.R.S., laid before the Meeting the neck-
vertebre of a young male Giraffe (Camelopardalis giraffa) which
had died in the Society’s Gardens on Jan. 8th, and made the
following remarks :—
It will be remembered that this animal in life showed a per-
manent bend in the neck, which was slight and hardly noticeable
at the time of its arrival, but increased greatly before the time of
its death. After death the neck-vertebre were carefully cleaned
and have revealed the causes of this bend, which undoubtedly
pressed upon the spinal cord. There was no tumour of any kind,
bony or otherwise, the existence of which might possibly have
been presupposed from the external appearance of the neck.
The bend in the neck was in fact related to the following condition
of the cervical vertebre.
The vertebre chiefly affected—but, as will be seen presently,
not the only ones affected—are the fourth and fifth. These two
vertebre are in the first place firmly ankylosed together so as to
be perfectly immovable the one upon the other. The bend
occurs in this region, and is produced by these two vertebree which
lie in relation to each other at an angle of nearly 90°. This bend
is due to an overgrowth on one side of these vertebre, the left,
and a cessation of growth on the other side. This overgrowth
mainly concerns, so far as I can make out, the epiphyses
of the vertebre in question. The general appearance produced
is that both vertebre are shorter in relation to the adjacent
vertebre than the normal. I have observed that the fourth and
fifth vertebre are the two which have been mainly affected. Of
these the fourth is more altered than the fifth. The neural
spine of the fourth vertebra is curved towards the left in relation
to the curvature of the whole vertebra; that is to say, the
convex border of the curve is on the left side. In addition to
this the spine itself is bent over to the opposite side, 7. e. to the
right, and forms a cavity deep enough to hide the first finger.
Such a bending of the vertebral spine does not occur in the case
of the fifth vertebra.
It is interesting to notice that the adjacent vertebre have
made an attempt, so to speak, to rectify the curvature caused by
the injury to the fourth and fifth vertebre. This state of affairs
is naturally seen in the most marked degree in the two vertebree
immediately adjacent to those which have been injured.
Particularly is this the case with the third vertebra. This
vertebra is bent, but in the opposite direction to the fourth; it
is the left side which is concave. The spine too is curved in the
same direction, and there is a slight concavity formed in the same
way by a bending over of the spine. This, however, lies on the
left, side and not on the right as is the case with the fourth
vertebra. Even the axis vertebra is slightly asymmetrical, and a
1902. | MR. F. E. BEDDARD ON A GIRAFFE. 53
Text-fig. 9.
Cervical vertebrze of a Giraffe.
Neck, showing cervical vertebrae in situ; dorsal aspect.
At., atlas; B, overgrowth of fifth vertebra; C. vIt., seventh cervical vertebra.
54. PROF. W. B, BENHAM ON THE OSTEOLOGY [Feb. 4,
careful examination of the posterior half of the atlas shows that
it is not perfectly symmetrical. The sixth vertebra is distinctly
asymmetrical, but the seventh has retained its normal symmetry.
The drawing exhibited (text-fig. 9, p. 53) illustrates the facts
that have been dealt with.
Dr. Chalmers Mitchell, F.Z.S., read, on behalf of Mr. EH. Degen,
a paper entitled “‘ Ecdysis, as Morphological Evidence of the
original Tetradactyle Feathering of the Bird’s Fore-limb, based
specially on the Perennial Moult of Gymnorhina tibicen.” The
material on which the paper was based consisted of a large series
of specimens of the Gymnorhina obtained at regular intervals
throughout the moulting-period, and the author had thus been
able to give a very complete account of the perennial replacement
of the feathers, avoiding the errors due to observations on the
altered habits as produced by captivity. The author showed that
the moulting of the wing-feathers took place in definite groups,
and indicated a composite origin of the modern feathering. He
thought that the new facts brought forward strengthened his
already published theory of the wing-feathers being derived from
the feathers of a four-fingered manus. Incidentally he suggested
that the eutaxy of the Passeres was essentially different from that
of such primitive birds as the Galline.
This Memoir will be published in full in the Society's
‘ Transactions.’
The following papers were read :—
1. Notes on the Osteology of the Short-nosed Sperm-Whale.
By W. Buaxtanp Bznnam, D.Sc., M.A., F.ZS.,
Professor of Biology in the University of Otago, New
Zealand.
[Received November 8, 1901. ]
(Plates II.-IV.")
A specimen of the Short-nosed Sperm-Whale (Cogia breviceps) |
came into my possession in 1900, and I have already communicated
to the Society some remarks on certain of the viscera*. I now
wish to offer some notes on the skeleton.
The animal, a male measuring 8 ft. 9 inches, had been cast ashore
on the sandy beach at Parakanui, Otago; and though it_had been
a good deal cut about, I was able to obtain the entire_skeleton,
together. with the cartilaginous portions of such structures as the
hyoid, sternum, and limbs: these were put through the gelatino-
glycerine process without any previous separation from the bones,
1 For explanation of the Plates, see p. 62.
2 See P. Z. 8. 1901, vol. ii. p. 107.
IPA. Ss, IO a) oll Lei,
Parker & West imp.
MP. Parker lith.
OS PBOLOEG On CO Gila.
dey An Ss I Ore rs syaolla, VE JUNE,
~~
MP. Parker lith \ Parker d& West imp.
OSLBOLOEGY OF COGIA.
IPL AS), Wea, rol Ie NG
MP. Parker lith. Parker d West imp.
OSTHOLOGY OF COGIA:
1902. ] OF THE SHORT-NOSED SPERM-WHALE. 55
and are thus preserved in a natural condition, There is one bone
upon which some doubt must still be expressed, viz. the pelvis.
Wall (1) describes and figures this structure as consisting of two
pairs of more or less circular or oval plate-like bones, which he
arranges in a transverse row—an inner smaller and an outer
larger bone on each side; the bones are very unlike the pelvic
bones of other Odontocetes, and as they were found in the sand,
it is within the bounds of possibility that the identification is
incorrect,
I searched the Parakanui carcase carefully for the pelvis: I
removed the penis and found no bone in connection with it, and
I feel quite certain that no bone existed, for the maceration was
most carefully carried out, and the contents of the macerating-tube
were sifted, so that even the cartilaginous epiphyses of the larger
ribs were recovered; if there had been bones of the size and
shape described by Wall, they could not have been overlooked.
The Axial Skeleton.
The total length of the dried skeleton, when the bones were
laid out, in contact, is 2°39 metres (7. e. 7 ft. 114 inches), of which
the skull measures 0°39 m. (153 inches) and the vertebral column
2°00 m. (6 ft. 8 inches). These measurements do not allow for
the intervertebral discs. I have not deemed it necessary to give
an account of the skull, as it has been adequately described and
figured by Owen (2), and more recently by Beneden & Gervais (5).
There is, however, one point to which I will refer, as it seems to
have escaped the notice of previous authors.
At the tip of each premaxilla is a short triangular calcification—
apparently not bone, but calcified cartilage, for it differs consider-
ably from bone, both in colour and texture (Pl. II. fig. 1, X).
Each of these ‘“sclerites,” or premaxillary nodules as they may
be termed, is grooved along its lower surface, and in this groove
lay the base of the single tooth of the upper jaw. ‘This groove is
in line with that on the maxillary bone, which is continued
backwards as a canal, to join the infra-orbital canal.
The premaxillary nodule is not indicated in Owen’s figure, in
which the upper tooth is placed in the anterior end of the
maxillary groove, and not on the premaxilla at all,
I have not seen the figure given in Van Beneden & Gervais’s
work, but no mention of the nodule occurs in the text : indeed,
these authors express some doubt as to the existence of the upper
teeth (p. 349). In a second skull in the Dunedin Museum,
belonging to an older specimen, obtained from Napier, in the
North Island, this premaxillary nodule does not exist; nor is
there any sign that it has fused with the premaxillary bone, for
the form of the latter and its relations to the maxilla are precisely
the same as in the Parakanui skull, if the nodule be removed.
No doubt this nodule remains separable from the bone, and hence
the absence of the upper teeth in most of the skulls of Cogia.
56 PROF. W. B. BENHAM ON THE OSTEOLOGY [ Feb. 4,
In the lower jaw of my specimen there are 13 teeth on each
side; but in the Napier skull I find 15. The former is the
number given by Wall and Krefft (3); the latter number is
attributed to this whale by de Blainville and by Van Beneden &
Gervais; while Flower & Lydekker, in their text-book on the
Mammalia (first edition), give the number as from “9 to 12.”
The Vertebral Column.
Turning to the vertebral column, there are, in addition to the
cervical mass of seven fused vertebre, in which all trace of
the separate vertebre is absent, except of the Ist, 2nd, and (th,
46 free vertebre, of which 13 are thoracic, 9 are lumbar, 23 are
caudal, of which the anterior 13 bear chevron-bones. In this
enumeration I have followed Flower (on Physeter) in regarding
as the first caudal that vertebra which carries at its hinder end
the first chevron.
It may be useful to give a summary of the formula of the free
vertebre according to previous authors’.
aoe Mo: of Thoracic. | Lumbar.| Caudal. | Chevrons.
| |
IVa eevee asec 44 14 9 21 | 13
Von Haast... A3 12 Thal 20 | 8
iKeretitee. cone: 48 13 9 20 eo
v. Beneden | | 2 | = oiellameaea
& Gervais } : ; 48 Sane: | :
With regard to the number of thoracic vertebre, there thus
appears from the accounts to be some slight discrepancy. Von
Haast has already pointed out that the total number of vertebre
“52.” given by Wall, is due to an error in addition of the
constituent vertebre. Van Beneden & Gervais state (p. 351) that
“whereas Wall describes 14 thoracic vertebre and 14 pairs of
ribs *, we only count 13 on our figure.” They suggest that perhaps
the small 14th rib, being free and independent of the vertebral
column, had disappeared during the preparation of the skeleton
(C. macleayi), and they found only 13 thoracics in the Japanese
specimens described on p. 515.
In the Parakanui skeleton there are only 12 pairs of complete
ribs articulated with the vertebre; but amongst the debris of the
macerating-pan I found a small bone (Pl. II. figs. 2, 3),
measuring only 37 mm. in length by 9°5 mm. in greatest breadth.
1 Flower & Lydekker give C. 7; Th.18 or 14; L.+C. 30: total 50 or 51.
2 Wall found only the ribs of the right side.
1902.] OF THE SHORT-NOSED SPERM-WHALE. 57
The two ends are rough and evidently had cartilaginous con-
tinuations. One end is broader than the other, and is apparently
the lower extremity : one surface is flat, and this I take to be
the external surface; the other is very convex from side to side
(see Pl. II. fig. 2, @), meeting the flat surface in a more or less
sharp edge; one edge, the anterior, being much sharper than the
other.
The general form of this little bone agrees very closely with
the shape of that region of the 12th rib just distal of the
curvature; here the outer surface is flat and the inner surface
convex, the outline of a transverse section being (as shown in
Pl. II. fig. 2, a) similar to that of the above small bone.
Further, I discovered a narrow, curved cartilage, four inches
(100 mm.) in length, pointed at one end, truncated and slightly
excavated at the broader end, which fitted on to the broad end of
the small bone. There is no doubt in my mind but that this
bone and cartilage constitute part, and the greater part, of the
13th rib of the left side ; the upper end of which must have been
connected to the 13th thoracic vertebra: the connection was
probably by means of cartilage, for this narrower upper end of
the bone is rough and convex.
On re-examining this vertebra, I noticed that the end of the
transverse process is similar to that of the 12th, and unlike that
of the succeeding vertebra, in that it hasa small articular surface
on the left side, but none on the right side.
We have here, I think, an explanation of the discrepancy as to
the number of thoracic vertebra; for, except in a very carefully
macerated skeleton, this little bone would undoubtedly be over-
looked ; and in skeletons lying on the shore there is little
likelihood of this last rib being found. Wall’s figure, however,
is erroneous in that he places the last rib (the 14th according to
his enumeration) in line with the lower end of the preceding ;
but from the form of the bone and its resemblance to that part of
the preceding rib, I think that it lay higher up, in the position
indicated in PI. II. fig. 2, with a long strip of cartilage below,
and a shorter cartilage (which I did not succeed in recovering)
above.
In Wall’s specimen this last small rib measured 13 inches, and
the preceding rib 114 inches. Krefft, too, notes that the last
rib, the 13th, is but 4 inches in length, whilst the preceding is
12 inches. It is not stated whether the measurement of the rib
was taken along the curve, or in a straight line from the capitulum
to the free end, but presumably it was in the former manner.
In my specimen the 12th rib is 9°6 inches (235 mm.) along
the outer curve, or in a straight line 83 inches (215 mm.), and
the bony part of the 13th rib is 14 inches (37 mm.). I estimate
that the total length of this rib, with both upper and lower
cartilages, was about 8 inches (200 mm.).
We may then conclude that in Cogia there are 13 thoracic
vertebree, with 12 pairs of complete ribs articulating with the
s
98 PROF. W. B. BENHAM ON THE OSTEOLOGY | Feb. 4,
column, and that the last (13th) rib is imperfectly ossified and
that the bone does not reach up to its vertebra, except possibly
in very old individuals.
The Sternum.
As far as I have been able to discover, the sternum of this
whale has not yet received an adequate description. It was only
partially recovered by Wall, who gives but a short account of the
imperfect bone, while it is not referred to either by von Haast (4)
nor by Van Beneden & Gervais.
In the Parakanui specimen the sternum (PI. III. fig. 4)
consists of three sternebre; the first and second formed of a
single bone apiece, the last of a pair of small bones. Hach sternebra
is capped by cartilage at each end, and the posterior end is bifid.
The anterior end of the sternum is bent slightly upwards, but
otherwise the bones are flat; the thickness increases from the
anterior end, where it is 8 mm., to the hinder end, which is
13 mm. in depth. The first two sternebre have rounded lateral
margins, while this margin, in the case of last pair of bones, is an
abrupt slope downwards and outwards from the dorsal surface,
with a sharp but obtuse upper and a sharper acute lower edge—
the ventral surface of this last sternebra being wider than the
dorsal surface.
There are four cartilaginous sternal ribs, measuring 90, 75, 60,
and 30 mm. respectively.
The following measurements were made :—
miliim.
Total length, including cartilage.....................06- 260
Greatestibresd thy. kel tits Meena aman see cattenimecs 155
Weastibreadtin 7.2 ONt, Ae CAN, oie SR ne 45
Length of the first bony sternebra* along the lower
SUTAcent, EN ra we ene cee Ree einen eee 90
Greatestibreadtht Ok Ie, it a Ne Le Le al eee 100
iBreadthvat posterionrendis..ch rere ener eee ence eee ek 60
Thickness (dorso-ventrally) in middle.................. 10
Length of second bony sternebra ..............0ee ee 76
breadth at anterior Onde funwecsessescepeciae meee: 54
Get CMMI CULC Se athe cana eae Raw ee ni tana en naar oee 43
wa Rab MOSLELIOL CRO iiss. cace acon aehcere cece ite 51
AMAT CEM OSSIAN OER cat ce sere sderaune dee team cme a tcate deste 12
Length of each ossicle of the 3rd sternebra ......... 3
Greatest biteadtii' |, ORE es ONS sin con nnce eee meea ts 20
MWe Kia CSS ie Pet Ee eA ees tase tenet ae aes ete oe 13
The Hyord.
The hyoid is very briefly referred to by Wall, and rather more
fully described (with a figure) by Van Beneden & Gervais.
1 Since the cartilages are only exceptionally preserved, the measurements of the
bones are also given.
1902.] OF THE SHORT-NOSED SPERM-WHALE. 59
In the present specimen (see Pl. III. fig. 5) the bones and
cartilage were uninjured.
The basihyal is a flat, irregularly circular bone, notched in the
middle line posteriorly, and with a pair of slight prominences at
the anterior end, separated only by a shallow furrow; each of
these prominences bears a small tetrahedral cartilage, which
evidently correspond to the bony projections seen in Physeter, but
which in Cogia do not appear to ossify, for they are unrepresented
in V. Beneden’s figure.
The anterior cornu consists of two segments, viz.: a short
proximal, curved cartilage, circular in section, representing the
ceratohyal; and a much longer distal region, the middle of which
ossifies to form the cylindrical stylohyal bone. The posterior
cornu, as in Physeter, is a broad plate of cartilage, in the midst
of which is a more or less circular flat bone—the thyrohyal bone.
This posterior cornu is not segmented from the basihyal, the
cartilage being perfectly continuous.
Measurements.
Basihyal bone : -millim.
Crreatest toread tiny se ysciase. esmeitcron danienice ccln/esairiete a seach 84
. eTNCAH 0 Bae Gao nonaus sdeboe ouAbes yo sebaconudeetooeone: 66
INBIIGSAVESIS. Gdodne sa se58 oe Saab bOOt dono odoSsoaDuReoosapEone apnond 5
Length of cartilaginous process ............sceesseeseneees 18
Total length of each half of the basihyal + thyro-
hyal, from the anterior end of the cartilaginous
PLCCESS) LOMDIp Ol COMMU, cle saReree lsc scr: HiAtioces 156
Greatest breadth, across the two posterior cornua,
measured from the outer margins ...............06 188
Wenge throtepmyroliyal bone ween seme deca acces ce 55
Breadth of ig Wn eA fate stil re tamnedoieae aie ae eee aat 46
Amberioricormuy total lengths. erence passe cacme cee 220
Length of ceratohyal cartilage (in middle line)......... 37
: Stilohiyali segment at uanseaepiisccresaeds ccc 175
. 5 bone (along its middle)............ 65
a ae hind ery Maren See esc s.. aesssaeile (65)
IMMIGESTIESS aE dooanobsehoos sobqdsnoboogbonboddes sbangobonsuades0 15
The Scapula.
This bone has been figured more or less accurately by all the
authors who have dealt with this whale, but without the cartilages.
The scapula has the usual cetacean form (see Pl. IV. fig. 6); its
external surface is feebly concave, owing to the reversion of the
anterior margin and of the superior border. The spine is but
feebly developed, but the acromion is a large subquadrangular
process. The coracoid process is large and well marked, not quite
so long as the acromion. The glenoid cup is oval.
The following measurements of the bone, without its cartilage,
were made :—
60 PROF. W. B. BENHAM ON THE OSTEOLOGY [ Feb. 4,
millim.
Greatest height (from the highest point of the
superior border to the anterior margin of the
POM OIL) iss ca Rasaiys oe AN Gas ema eeeR ne Gas Rie eee 164
length of the posterior border ......................-. 107
Be ANLCRION Cord VR nite et Ake eee 159
Greatest breadth (in a straight line from anterior
to posterior angle of the superior border) ...... 184
Breadth immediately above acromion ............... 3
wength ot elenoid Cupp yee eeneeneine see en ee 46
Breadth es wy uyesterceras ce Tyas it ci Bee sectcerannamastnneit 31
Distance from the antero-superior angle to origin
OlpaerOMMONs Maem bas. Sar fuk leakiscos teceeee oe ik 76
eniethyonacromlomen ncaa. a. canna ieee eeEe Ren ean eee 48
Vertical height (near root) of acromion............... 35
Distance from posterior margin of glenoid to tip of
ACVOMMON “Ren sovdnas teases wn eaeanenae enon tees 101
Length of coracoid (from anterior margin of glenoid) 47
Distance from posterior margin of glenoid to end
OL COrACOIG ante ene eee aie ae ar tecae enon 84
The Pectoral Limb.
The limb has been more or less imperfectly figured by the
various authors—a photograph of the “restored” limb having
been added to the second edition of Wall’s memoir, to replace an
inaccuracy in the figure of the entire skeleton.
In this photograph, the restored carpals (which were gathered
from the sand and pieced together) are fairly accurately placed ;
but the cartilages, having been represented by some artificial
filling, do not show their characteristic independence. Wall
describes “seven” carpals, but it is evident from later researches
that the “two linear transverse bones” are merely the distal
epiphyses of the radius and ulna, at the ends of which he locates
them; the remaining 5 are accurately described in the text. The
photograph is a truer representation of the hand than the woodcut
accompanying Krefft’s paper.
The figure given by Van Beneden & Gervais is also incomplete.
It seems therefore worth while to present a complete figure of
the entire limb (Pl. IV. fig. 7) showing all the cartilages and
bones in their true position.
The humerus is provided with a small deltoid ridge, 15 mm.
in length and 5 mm. in height. The head and tubercle, as well
as the distal epiphysis, are embedded in cartilage, but are firmly
united to the shaft of the bone. But the epiphyses of the radius
and ulna are not as yet united, though they can be felt at each
end by a needle thrust into the cartilage.
The proximal epiphysial cartilage of the ulna is prolonged down-
wards as a spur, which represents the bony olecranon of Physeter.
This cartilage is indicated in the figure given by Krefft, and in
the photograph of Wall’s specimen, as a small bony process. In
my specimen there is no ossification in this cartilaginous olecranon.
1902. ] OF THE SHORT-NOSED SPERM-WHALE, 61
The distal epiphysial cartilage of the radius is produced along the
outer sides of the carpus up to the metacarpal of the first digit,
so as almost to suggest a carpal; but as each of the true carpals
has its own cartilage around it, this prolongation seems to have
some other significance.
The carpal bones are five in number, three belonging to the
proximal row, and two to the distal series. Each is an irregular
polygonal, more or less hexagonal, disc of bone embedded in
its own cartilage. Hach bone has vertical sides, without the
“shelf” and without the epiphysis which exist in Physeter, to
which, otherwise, they bear considerable resemblance. The
pisiform is entirely cartilaginous. In the digits, each phalanx
is provided with its own independent cartilaginous epiphysis at
each end, as in Odontocetes generally. The metacarpals are
short, not much longer than the proximal phalanx in each digit.
That of the first digit is, as in Physeter, rounded and somewhat
like a carpal; but Flower has given reasons for regarding this
as a metacarpal, and the fifth metacarpal is also rounded. The
relative lengths of the digits, in ascending order, are I., V., IV.,
JMS, Is
The number of phalanges is accurately shown in the drawing ;
whereas in the previously published figures some of the terminal,
very small phalanges are missing.
The first digit possesses two phalanges ;
i second x ss ten ‘3
a third.) 5 by seven ,,
ee LOUt Ia ee Six Ee
eee tint amare ss three _,,
These numbers refer to the right limb; on the left the second
digit has only nine and the fifth only two phalanges.
It will be noted that the two to four terminal phalanges of the
longer digits are more or less circular, as are all three of the fifth
digit.
Measurements.
millim.
Motalblemert loge coccos dct stultvadins «ve Bban Thlesicmndeosindiciisiamtnseanpas 372
Humerus :—Length (incl. cartilage) .................:00000 95
Pee sha itromllveie wicca ce eee eee 65
Transverse diameter of shaft at WPPeRVeN Cape. ceee ees 45
3 lowerrendincereene er 50
Girth, inlite middle: «MWA eee vail ae 101
ARR CIWVESS) LGR WAGs Me tO EY Cel a eee aah te kta ies) 24
Radius :—Length along preaxial border .................. 75
“A sr) POstamaal border ay qd qececccs nc 60
», of bone only (in middle line) ......... 60
lheast orveadthi 05: ieer cn eeecenenetr etch ts veins cence 30
is or) DIAL CAN OSS) sai est Seen Aen Rabe ne lateab aise asics ecole 12
Ulna :—Total length along postaxial border ............... 63
6 o oy. reamal Moonder: sce. ceniesn 60
ay oe Moisl corn) (aT EIS): Ge Us banwacawsmabones 55
TEAS) ATCC ee en one oh li 26
Gila. leg OR GI ga:cia toa ora ores Ce eertar Eicon «$0
2
62 ON THE OSTEOLOGY OF THE SHORT-NOSED SPERM-WHALE. [ Feb. 4,
millim
Olecranon, length .........cccsceeeneeeeeee ones Be ancinaaite setae qe 25
Total breadth of carpus ........+.++. MPL AR NC aero ance 80
Digits: Total length, including cartilage :-—
Meriphand:-s0 00; (digihes cece secsea eee Sstosa ns elaine stesiSis 52
5 TE pi ee acre sehr iete ase eie sean aoa 185
* DD i os vse wsemaeecnenteas ede Roseecee 158
Bs AV es a aster ceca senigeees saiscaieeeewsies 114
- WV sain ua hoses sina Jue CORE aaar oat Opies 52
Right hand ¥2 Fo 5, .--c00+5.5-- Seige sicelc Semin se seine seis at 55
is 1 few BRS See apis Meageeh esc asenancte lOc
- Ag pi gla ee cise atetaatas Sena eaedacee 148
a TENE tee ar sel ec Se 102
Vic 68
List of the memoirs to which references are made.
1. Wat.: “ History and Description of the Skeleton of a New
Sperm-Whale.” Sydney, 1851 (reprinted 1887).:
. OwEN: “Onsome Indian Cetacea.” Trans. Zool. Soc. vi. 1865,
p. 30.
: “ Notice of a New Species of Sperm-Whale.” Proce.
Zool. Soc. 1865, p. 708.
_ y. Haast: “On the Occurrence of a New Species of Huphysetes
on the Coast of New Zealand.” ‘Tr. N. Z. Institute, vi.
1873, p. 97.
5. Van BeneDEN & Gervais: ‘Ostéographie des Cétacés,’ pp. 349,
515, pl. 61 (1880).
m ow »
a
SI
rj
ty
A
EXPLANATION OF THE PLATES.
Puate II.
Fig. 1. Anterior end of the ventral surface of the skull of Cogia breviceps (X 4),
showing the paired premaxillary nodules (X) carrying the teeth (¢).
gr., maxillary groove. m., maxilla. pmzx., premaxilla. vo., vomer.
2. ‘The external surface of the last two ribs (xX #) showing what is believed to
be the true position of the rudimentary (13th) rib in relation to the 12th.
The cartilaginous lower end of the rib is dotted ; the upper region—indicated
by dotted outline—is the presumed continuation of the rib to its articulation
with the vertebra. At the side of each rib is shown the outline of its
transverse section (a).
3. View of the anterior side of the 13th rib. X 1.
Puate III.
. Sternum of Cogia breviceps, with the sternal ribs, dorsal aspect. X 3.
. Hyoid, of which only the right anterior cornu is represented, dorsal
aspect. X 3.
ic
bee
09
oe
Puate IV.
Fig. 6. The right scapula, external surface. X 3.
7. The right pectoral limb, external surface. Some of the distal cartilages
have been inserted from the more perfect left limb. X .
1 Some of the terminal cartilages were imperfect on this hand, and have been
restored in gelatine, but not quite accurately.
PZ.S. 1902, vol. 1.PLV.
Hy
"f
ES
gp
c
mo
sai
>
| ttf WN
pea vaieaaan Gays
\
en
ros)
sa
SS
SE reread ee ree
ice)
(cr i Lament a es
Edwin Wilson,Cambridge .
S FROM THE MALAY PENINSULA.
Et
Cy
DRAGONFLII
PZS1002) vol Sev
4
La Ege
iy
we
is
\\
~
anes eO=
i
eS ee
Vy
bridge.
.Cam
Nilson
rin V
Edw
1902.]| ON THE DRAGONFLIES OF THE “ SKEAT EXPEDITION.” 63
2. On a Collection of Dragonflies made by Members of the .
Skeat Expedition in the Malay Peninsula in 1899-1900.
By F. F. Larpiaw, B.A.
[Received December 28, 1901. ]
(Plates V. & VI." and Text-figures 10-12.)
In drawing up an account of the Dragonflies collected by
members of the Skeat Expedition, I have thought it worth while
to include in my list not only the names of species represented in
this collection, but also of all those which I have been able to find
recorded as having occurred in the Malay Peninsula. It will be
seen that the list is a fairly large one, although it is impossible to
suppose that the full richness of the fauna of this part of the
world has been as yet revealed.
Noticeably this is the case with the Gomphine ; it is worthy of
remark that our collection contained five specimens representing
four different species, and that none of these were identical with
any species previously found in the Peninsula. I have been able
through the courtesy of Mr. Kirby to add to my list the names
of the species taken by Mr. Ridley, specimens of which are in
the British Museum. I have to thank both Mr. Kirby and
Dr. Sharp very sincerely for many useful suggestions and much
kind assistance.
Lastly, I have to thank the other members of the Expedition
for their kind assistance in making the collection.
I have given references in every case where possible to Mr.
Kirby’s ‘Catalogue of the Odonata,’ published in 1890, where
full allusion to papers published before that date will be found.
The following notes on the habits of some of the species col-
lected may be of interest :—
LipeLLuLip#.— Almost without exception the numerous mem-
bers of this family avoid forests and are to be found in flat open
country, rice-fields, and clearings near the forests, especially where
there happens to be a stagnant pool in the neighbourhood,
Certain very common and widely spread species are to be
found wherever there is a suitable locality. Such are especially
Orthetrum sabina and to a lesser extent Pantala flavescens,
Tholymis tillarga, Trithemis trivialis, Trithemis aurora.
Certain other species with a very wide range in the Oriental
Tropics seem to prefer the neighbourhood of the sea. Such are
the members of the genus Ryothemis, also Neurothemis tullia and
Brachythemis contanvinata.
The rarer and more characteristic species are only to be found
in up-country clearings. The only species that I saw actually in
the forests were Camacinia gigantea, Cratilla metallica, Tyrio-
1 For explanation of the Plates, see p. 92.
64 MR, F, F, LAIDLAW ON THE | Feb. 4,
bapta torrida, Orthetrum pruinosum, and Calothemis biappendi-
_culata. Others for the most part were caught playing round
stagnant water. Rapidly running streams are invariably avoided
except by Zyriobapta torrida.
On the other hand, the AiscHNIDZ are mostly found in the
forests, any small stagnant pool is an excellent locality; the
species of Gynacantha and Anax guttatus are sometimes seen in
the open. The large species of the Gomphine are also forest
insects. Thus my specimen of Sieboldius grandis was taken in
the same locality (a small muddy pool frequented by wild pig)
with two males of Amphieschna ampla; this locality also yielded
Pericnemis stictica and Lestes ridley.
Another forest-haunting group is found amongst the Cato-
PTERYGIDZ. Vestalis amena never occurs in the open, nor over
rapidly running water: probably Hecho and Climacobasis have
similar habits ; they resemble Vestalis amcmna so closely that they
may perhaps be often mistaken for this very common species.
The other Calopterygine are only to be found playing over rapidly
running streams and rivers, and their beautiful iridescent wings
add greatly to the charms of a sun-lit river-scene. Rhinocypha
fenestrella sometimes forsakes the main stream for the shady
rivulets that wander through the forest, but most of the species
prefer the wider waters. The lovely Vewrobasis chinensis wanders
farther down the river perhaps than other species, but I have
never seen it near the mouth of a river, or in fact after the
stream had become sluggish and polluted.
Of the Agrionine numerous species are found in rice-swamps :
few make their home in the forests, amongst these are Pericnemis
stictica and Lestes ridleyi referred to above, as well as one or two
species of Psilocnemis, Amphilestes, and a few of the Protoneurous
group.
In many genera the females are exceedingly rare; this is
especially the case with the Calopterygine genera Huphca and
Dysphea. It has been suggested that the soberly coloured
females do not attract the notice of collectors to the same extent
as the males, and that hence they are rarely found in collections :
but I can assert positively that in their own haunts the females
are exceedingly rare; to the best of my belief, I saw only one, a
female of Huphea ochracea, which I secured.
Family LIBELLULID.
Subfamily LIBELLULINA.
(Species marked with an asterisk are not represented in our Collection.)
*ZYXOMMA PETIOLATUM Ramb.
Zyxommea petiolatum, Kirby, Cat. Odonata, p. 35.
East Indies. Singapore (/idley).
1902. ] DRAGONFLIES OF THE “ SKEAT EXPEDITION,” 65
THOLYMIS TILLARGA (Fabr.).
Tholymis tillarga, Kirby, Cat. Odonata, p. 1; Selys, Ann. Mus.
Genov. (2) x. p. 439.
Common in the Eastern Tropics.
PANTALA FLAVESCENS (Fabr.).
Pantala flavescens, Kirby, Cat. Odonata, p.1; Selys, Ann. Mus.
Genoy. (2) x. p. 440; Ris, Arch. f. Naturg. Jahrg. 66, p. 175.
Found in the tropics of both worlds.
ae
CAMACINIA GIGANTEA (Brauer).
Camacinia gigantea, Kirby, Cat. Odonata, p. 2.
Two fine males were taken at Kwala Aring, where this species
is fairly abundant near pools in open spaces. It is very ditlicult
to catch, being a powerful flier. It haunted the same localities
as Neurothemis stigmatizans, which resembles it very closely in
colour, though of course much smaller.
HyYDROBASILEUS EXTRANEUS (Hagen).
Hydrobasileus extraneus, Kirby, J. Linn. Soc., Zool, xxiv.
PLol’, ple xditig Moe
Recorded from Penang.
RHYOTHEMIS PHYLLIS (Sulz.).
Rhyothemis phyllis, Kirby, Cat. Odonata, p.5; id. Journ. Linn,
Soc., Zool. xxiv. p. 549; Selys, Ann. Mus. Genoy. (2) x. p. 443.
This species is common along the east coast of the Peninsula.
Specimens were collected at Singgora, Kota Bharu, Kelantan, and
at Trengganu. Occurs throughout the Malay Archipelago.
*RHAYOTHEMIS FULGENS Selys.
Rhyothemis fulgens, Kirby, Cat. Odonata, p. 6.
Singapore (Selys); Dindings (Ridley). Borneo, Malay Penin-
sula, Sumatra.
* RHYOTHEMIS CURIOSA Selys.
Rhyothemis curiosa, Kirby, Cat. Odonata, p. 6.
Singapore (Selys). Sumatra. Perhaps a race of FR. fulgens
(Selys, Ann. Mus. Gen. xxvii, p. 451).
* NEUROTHEMIS FULVIA Drury.
Neurothemis fulvia, Kirby, Cat. Odonata, p. 7.
Neurothemis sophronia, Selys, Ann. Mus. Genov. xiy, (1879)
p. 292.
Malacca (Selys). China, Bengal, Nepaul.
NEUROTHEMIS FLUCTUANS (Fabr.).
Neurothemis fluctuans, Kirby, Cat. Odonata, p. 7; Selys, Ann.
Proc. Zoou. Soc.—1902, Vou. I, No. VY. 5
66 MR. F. F, LAIDLAW ON THE [| Feb. 4,
Mus. Genov. (2) x. p. 446; Karsch, Abh. v. d. Senckenberg. nat.
Gesell. xxv. 1. p. 219.
Common at Kwala Aring. Widely spread in the Eastern
Tropics.
NEUROTHEMIS STIGMATIZANS (Fabr.).
Neurothemis stigmatizans, Kirby, Cat. Odonata, p. 7; Karsch,
Abh. v. d. Senckenberg. nat. Gesell. xxv. 1. p. 218.
Plentiful at Kwala Aring. Like the last a common and
variable insect.
NEUROTHEMIS DISPARILIS Kirby.
Neurothemis disparilis, Kirby, Cat. Odonata, p. 8.
Two specimens from Kwala Aring. Singapore (Ridley);
Borneo.
NEUROTHEMIS TULLIA (Dru.).
Neurothemis tullia, Kirby, Cat. Odonata, p. 8; id. Journ. Linn.
Soc., Zool. xxiv. p. 550.
Common near the mouth of the Kelantan River and for some
thirty miles up the river. A common Eastern species.
TRITHEMIS (?) TRIVIALIS (Ramb.).
Trithemis trivialis, Kirby, Cat. Odonata, p.18; id. Journ, Linn.
Soe., Zool. xxiv. p. 550 (1894).
Trithemis (2) trivialis, Selys, Ann. Mus. Genov. (2) x. p. 467
(1891); Kirby, Ann. & Mag. Nat. Hist. (7) v. p. 531 (1900).
Diplacodes trivialis, Karsch, Abh. v. d. Senckenberg. nat.
Gesell. xxv. 1. p. 219.
Widely distributed, ranging from India and Ceylon to Japan.
I obtained specimens at Kwala Aring and Kota Bharu, Kelantan.
Taken also by Mr, Ridley in Province Wellesley.
As pointed out by Mr. Kirby (Ann. & Mag. loc. cit.), this
species probably requires the creation of a new genus to receive it.
TRITHEMIS AURORA (Burm.).
Trithemis aurora, Brauer, Verh. zool.-bot. Ges. Wien, xviii.
p. 117 (1868); Selys, Ann. Mus. Genoyv. (2) x. p. 465 (1891).
Trithemis intermedia, Kirby, Proc. Zool. Soc. 1886, p. 327,
pl. 33. fig. 4.
Trithemis yerburii, Kirby, Cat. Odonata, p. 18.
Trithemis aurora, Kirby, Journ. Linn. Soe., Zool. xxiv. p. 551.
This beautiful species was fairly common in September in
marshy rice-fields at Ulu Aring. Mr. Ridley has collected it
in Singapore.
BRACHYTHEMIS CONTAMINATA (Fabr.).
Brachythemis contaminata, Kirby, Cat. Odonata, p. 21; id.
1902. ] DRAGONFLIES OF THE “ SKEAT EXPEDITION.” 67
Journ. Linn. Soc., Zool. xxiv. p. 551; Selys, Ann. Mus. Genov.
(2) x. p. 468 (1891).
A widely spread Oriental species; common on the lower reaches
of the Kelantan River and in the town of Trengganu.
CrocoTHEMIS SERVILIA (Drury).
Crocothemis servilia, Kirby, Cat. Odonata, p. 21; Selys, Ann.
Mus. Genoy. (2) x. p. 468 (1891).
Kwala Aring in August, in an open space near forest. East
Indies and Australia.
BRACHYDIPLAX MARIA Selys.
Brachydiplax maria, Kirby, Cat. Odonata, p. 22.
Kwala Aring. Dindings and Selangor (Adley). Borneo.
*BRACHYDIPLAX .MELZNOPS Selys, Aun. Mus. Genoy. xxvil.
p. 457.
Brachydiplax melanops, Kirby, Cat. Odonata, p. 22.
A small species from Selangor taken by Mr. Ridley, and now
in the British Museum, probably belongs to the species indicated
by de Selys, agreeing with it in its small size. Abdomen 16°5 mm.
long; hind wing 22°5. The thorax and first fore segments of
abdomen blue-pruinose. 6 prenodals and 5 postnodals on the
fore wing. Internal triangle free.
BRACHYDIPLAX PRUINOSA, Sp. Nn.
Length of abdomen 18°5 mm. Length of hind wing 24 mm.
3. Head yellowish grey, margins of the upper and lower lips
black, frontal tubercle, and upper surfaces metallic blue. Eyes
brown.
Prothorax and thorax coppery green dusted over with very
pale blue ‘bloom.’ Abdomen: first five segments grey, also coated
with ‘bloom,’ the rest black, second and third segments with a
transverse carina; legs black; pterostigma and venation black.
Fore wings: 8 antenodals, 6 or 7 postnodals, Discoidal tri-
angle free, followed by two rows of cells.
Hind wings: 7 antenodals, 6 or 7 (usually 7) postnodals. The
hind wings have a faint tint of yellow at their base,
Two males from Kwala Aring taken in August.
* MICRODIPLAX DELICATULA Selys.
Microdiplax delicatula, Kirby, Cat. Odonata, p. 22.
MacropIpLaXx virttata Kirby.
Urothemis vittata Kirby, Journ. Linn. Soce., Zool. xxiv. p. 552,
pl. 42. fig. 2.
A male specimen from Kwala Aring. Mr. Kirby tells me that
this species should be referred rather to the genus Macrodiplax
5*
5
68 MR. F. F. LAIDLAW ON THE [Feb. 4,
than to Urothemis. The last postnodal cell is as long or a little
longer than the pterostigma in the fore wing.
TYRIOBAPTA TORRIDA Kirby.
Tyriobapta torrida, Kirby, Cat. Odonata, p. 32; Karsch, Abh.
v. d. Senckenberg. nat. Gesell. xxv. 1. p. 221 (1890).
This species haunted a small forest stream close to the village
of Kwala Aring. It was apparently confined to this locality in
that neighbourhood. A common Bornean insect.
CRATILLA METALLICA (Brauer).
Protorthemis metallica, Kirby, Oat. Odonata, p. 30; Selys, Ann.
Mus. Genov. (2) x. p. 461; Karsch, Abh. v. d. Senckenberg. nat.
Gesell. xxv. 1. p. 221.
Nesoxenia metallica, Kirby, Cat. Odonata, p. 180.
Cratilla metallica, id. Ann. & Mag. Nat. Hist. (7) v. p. 542.
Common at Kwala Aring and on Gunong Inas.
ORTHETRUM SABINA (IIl.).
Orthetrum sabina, Kirby, Cat. Odonata, p. 35.
Abundant all along the East Coast. Ranges through the Hast
Indies to Australia.
ORTHETRUM PRUINOSUM (Burm.).
Orthetrum pruinosum, Kirby, Cat. Odonata, p. 38; Ris’, Arch.
f. Naturg. Jahrg. 66, p. 185, pl. ix. fig. 3.
A single specimen (3) from Kwala Aring, September 1899.
East Indies.
ORTHETRUM TESTACEUM (Burm.).
Orthetrum testaceum, Kirby, Cat. Odonata, p. 39.
A pair, in cop., from Kwala Aring, September. Also a single
male from the same locality. Recorded from Java.
ORTHETRUM NICEVILLE Kirby.
Orthetrum nicevillei, Kirby, Ann. & Mag. Nat. Hist. (6) xiv.
p. 112 (1894).
Described from specimens from Tenasserim. A single specimen
from: Ulu Aring, September 1899.
OrTHETRUM. sp.—Our collection includes a female Orthetrwm
belonging to a species distinct from, but closely allied to, O. sabina.
The abdomen is shorter, 26 mm., and distinctly stouter, the anal
appendages are black, and the sides of the thorax are not so
distinctly marked with black. I have been unable to identify it.
LyRIOTHEMIS PRIAPEA Selys.
Lyriothemis priapea, Kirby, Cat. Odonata, p. 25.
This genus is closely allied to Orthetrum, but differs in the
1 Dr. Ris (loc. cit.) records O. chrysis from Malacca.
1902. | DRAGONFLIES OF THE ‘‘ SKEAT EXPEDITION.” 69
strongly curved sectors and in having three or four cross nervules
in the submedian space of fore and hind wings.
A single specimen, a male, from Kwala Aring.
PoTAMARCHA OBSCURA (Ramb.).
Potamarcha obscura, Kirby, Cat. Odonata, p. 180.
Potamarcha congener, Selys, Ann. Mus. Genov. (2) x. p. 459.
Potamarcha obscura, Karsch, Abh. v. d. Senckenberg. nat.
Gesell. xxv. 1. p. 219.
This species is common at Kwala Aring, where I took two
females and several males. Closely allied to Lathrecista, it differs
in having the eighth abdominal segment in the female dilated, and
the triangle of the hind wing traversed. (See also Selys, loc. cit.)
LATHRECISTA TERMINALIS Kirby.
Lathrecista terminalis, Kirby, Cat. Odonata, p. 30.
A single male from Kwala Aring. Recorded and described
from Borneo.
* LATHRECISTA SIMULANS (Selys).
Lathrecista simulans, Kirby, Cat. Odonata, p. 30; Selys, Ann.
Mus. Genoy. (2) x. p. 458.
Recorded from Borneo, Sumatra, Ceylon, Malacca, and Burmah.
* AGRIONOPTERA LINEATA Brauer.
Agrionoptera lineata, Kirby, Cat. Odonata, p. 31; Selys, Ann.
Mus. Genov. xix. (1879) p. 302.
Malacca. Philippines.
* AGRIONOPTERA MALACCENSIS Selys.
Agrionoptera malaccensis, Selys, Ann. Mus. Genov. xxvii.
p. 461; Kirby, Cat. Odonata, p. 31.
(This genus differs from the preceding in the absence of the
supernumerary antenodal nervule of the front wings, and in
having several cross nervules in the submedian space, as well as
in the position of the base of the triangle of the hind wings, in
front of the arculus. The two genera closely resemble each other
in coloration.)
* AGRIONOPTERA NICOBARICA Brauer.
Agrionoptera nicobarica, Kirby, Cat. Odonata, p. 31.
Singapore, Nicobar Is. :
* AGRIONOPTERA SEXLINEATA Selys.
Agrionoptera sexlineata, Kirby, Cat. Odonata, p. 31.
Recorded from Malacca.
*CALOTHEMIS BIVITTATA (Ramb.).
_ ~Calothemis bivittata, Kirby, Cat. Odonata, p. 42. -
70 MR. F. F, LAIDLAW ON THE [Feb. 4,
CALOTHEMIS BIAPPENDICULATUS Selys.
Calothemis biappendiculatus, Kirby, Oat. Odonata, p. 42.
6. Length of abdomen 22mm. Length of hind wing 28 mm.
Wings hyaline, slightly tinged with yellow at their bases.
Pterostigma black, 2 mm. in length, covering 3 cells.
Fore wing. 19 antenodals, the last continuous, 9-10 postnodals.
Discoidal triangle traversed, followed by two rows of cells. 2 supra-
triangular cross nervules, 2 cross nervules in the lower basal cell.
Internal triangle divided into 3 cells.
Hind wing. 2 supra-triangular, 3 lower basal cross nervules.
Discoidal triangle traversed.
Head. Lower lip yellow, upper lip black, eyes brown, occipital
triangle black, rest of head steely-blue black except a yellow mark
at the side behind each eye.
Prothorax black.
Thorax black above, dull brown below. Legs brown.
Abdomen. Segments 1 and 10 black, the rest bright red. Seg-
ments 2-3 with transverse carina. Segments 3-9 strongly
triangular in cross section, Rising from the bases of the genital
ramules are two long branches, standing at right angles to the
body.
2 unknown.
I took two specimens of this insect at Kwala Aring. They
differ from the type in having the upper surface of the thorax
rich black instead of brown. Otherwise they closely resemble it,
especially in the very remarkable genital organs on the second
abdominal segment.
*ORCHITHEMIS PULCHERRIMA Brauer.
Orchithemis pulcherrima, Kirby, Cat. Odonata, p. 42; Karsch,
Abh. v. d. Senckenberg. Nat. Gesell. xxv. 1. p. 228.
Singapore (Ridley). Malacca (Selys).
DIPLACODES NEBULOSA (Fabr.).
Diplacodes nebulosa, Kirby, Cat. Odonata, p. 42.
A single specimen was taken at Kota Bharu, Kelantan. There
are specimens in the British Museum taken by Ridley in Province
Wellesley. Widely distributed in the East Indies.
ACISOMA PANORPOIDES Ramb.
Acisoma panorpoides, Kirby, Cat. Odonata, p. 43.
Kwala Aring. One specimen, ¢. ‘Tropical regions of the
Old World.
TETRATHEMIS HYALINIA Kirby.
Tetrathemis hyalina, Kirby, Cat. Odonata, p. 44.
The two species of this genus which are represented in our
collection exhibit a very remarkable sexual dimorphism which
has not, I believe, previously been remarkesl. The males have a
1902. ] DRAGONFLIES OF THE ‘‘SKEAT EXPEDITION.” 71
very extraordinary development of the armature of the second
and third pairs of femurs. This development is paralleled in the
American genus Macrothemis and its allies and also in the Old
World genera Schizonyx, Neuwrocena, Zygonyx, and Zygonidia,
amongst other Lzbelluline (see Calvert, Pr. Ac. Philad. 1899,
p- 246).
eee hyalinia has in the male, on each of the second
pair of femurs, on their antero-inferior surface, 17 short straight
spines directed towards the knee, increasing gradually in size, but
the last three longer than the rest, more widely separated and
increasing rapidly. On the antero-inferior surface of each of the
third pair of femurs is a row of some 20 short curved teeth,
their apices directed away from the knee, decreasing gradually in
size distally; at the end of the series is a single short straight
spine directed towards the knee (see text-fig. 10).
Text-fig. 10.
Third femur of Tetrathemis hyalinia (X about 10).
TETRATHEMIS PULCHRA, sp. n. (Plate V. fig. 3.)
Length of abdomen, ¢ 16mm., 9? 15mm. Length of hind
wing, ¢ 17 mm., 2? 17°5 mm.
Wings hyaline, reticulation black. Fore wings tinged with
orange from the base about halfway to the nodus. Hind wings
tinged with orange about as far as the nodus.
Fore wings. 8-9 (usually 8) antenodals, 5 (in one case on one
side 6) postnodals; 1 supra-triangular cross nervule; 2 cross
nervules in the lower basal cell. Triangle followed by a single
row of cells.
Hind wings. 6 or 7 (usually 7) antenodals, 5 postnodals; no
supra-triangular cross nervule; 2 cross nervules in lower basal -
cell.
Coloration in the male. Face black with yellow marks as
follows: lateral lobes of lower lip, nasus, and rhinarium. The
vertex and tubercle are metallic coppery black. Back of head and
prothorax black.
Thorax black above, a few yellow spots between the wings.
Sides citron-yellow with two black bands. The first of these runs
from immediately in front of the first pair of wings obliquely
downwards to between the second and third pair of femurs. The
second runs from immediately in front of the second pair of wings
down behind the third pair of femurs. The whole ventral surface
is black, save that the yellow colour of the flanks extends for a
short distance over the ventral surface along either side The
legs are black, inner surface of first pair of femurs citron-yellow.
Abdomen black, with the following yellow marks :—a spot on
either side of segments 1-6, very small on 6, traversed by a black
72 MR. F. F. LAIDLAW ON THE [Feb. 4,
line following the transverse carina in 3-4. On segment 7a
dorsal yellow spot divided longitudinally by the black mid-dorsal
carina. Traces of a transverse carina are present on segment 5.
The abdomen is slightly dilated at its base, but from segment 4
onwards very slender.
Coloration in the female as in the male. The traces of a
transverse carina in segment 5 are more distinct laterally. The
abdomen is broader and of practically equal circumference
throughout.
In the male there are on the antero-inferior surface of the
femur 17 short curved teeth directed towards the knee, and
increasing in size distally very gradually. These are followed by
three straight spines inclined in the same direction; the first of
these is the shortest and the last the longest.
The third femur is provided on the antero-inferior surface with
a row of 23 thorn-like teeth with their apices directed away
from the knee. These increase gradually towards the distal end
of the femur.
*NANNOPHYA PyGMzA Ramb.
Nannophya pygmea, Kirby, Cat. Odonata, p. 45.
The British Museum has a number of specimens of this species
from Singapore.
Genera of uncertain position.
NEvROCENA IDA Hagen. (Plate V. fig. 1.)
Zygonyx ida, Hagen, Ver. Ges. Wien, xvii. p. 62; Brauer, op.
cit. xviii. p. 370 & p. 742; Selys, Ann. Soc. Ent. Belg. Alpe ws
id. Ann. & Mag. Nat. Hist. (4) ili. p. 274; id. C. R. Soc. Ent.
Belg. xxxv. p. CCXXVil.
Pseudomacromia luxuriosa, Karsch, Berl. ent. Zeitschr. xxxviil.
p. 21.
Zygonyx ida, id. Ent. Nachr. xxi. p. 203; Calvert, P. Ac.
Philad. 1899, p. 246.
Neurocena ida, Kirby, Ann. & Mag. Nat. Hist. (7) v. p. 541.
_ This appears to be an exceedingly variable species, and the
single specimen I obtained differs to a certain extent from those
described hitherto, so that it seems worth while to give a fairly
full account of it.
The length of the hind wing is 42 mm., of the abdomen 38mm.
The wings are hyaline, faintly tinged with yellow, which becomes
vivid towards the outer extremities of the fore wings. In a male
in the British Museum Collection the wings are almost colourless.
The reticulation is black.
Fore wings. 14 antenodals; on the left side the outermost is
continuous, on the right side discontinuous. 7 to 8 postnodals.
Internal triangle of both wings free, discoidal triangle free. (The
internal triangle is usually divided into two or three cells; de
Selys states that in two females examined the discoidal triangle
1902. ] DRAGONFLIES OF THE ‘‘ SKEAT EXPEDITION.” 3:
is free, in three females crossed; in males it is normally free.)
Two rows of post-triangular cells. Nodal sector strongly waved at
its middle. Arculus at the level of the second antenodal. Two
cross nervules in the submedian space.
Hind wings. 10-11 antenodals, 9-9 postnodals. Discoidal
triangle traversed. (According to de Selys the discoidal triangle
of the hind wing in the female is normally traversed; of 13 males
7 had it traversed and 6 free.) The triangle is followed by two
rows of cells. The British Museum specimen (male) has but one
row of post-trigonal cells in the hind wings. Pterostigma in the
females 1s about 3°75 mm. long, in the male about 2°25 mm.
Sectors of triangles of hind wings widely separated at their origins.
Rhinarium and nasus livid yellow. Frons and vertex metallic
blue-green. Thorax metallic blue-green. Abdomen slightly
thickened at its base, metallic black with fine transverse yellow
lines at the bases of segments 2 and 3 and on the transverse
carine of those segments. <A yellow spot on either side of the
second segment.
Legs black. In the males on each of the second pair of femurs
are a number of short teeth directed towards the knee. On the
third pair are 25 short teeth; of these the first eight or nine are
directed towards the knee, then follow one or two not inclined,
then eleven or twelve directed towards the trochanter, last one
or two not inclined. In the females all the teeth on the hinder
femurs are directed towards the knee.
ZYGONIDIA MALAYANA, Sp. D.
Length of abdomen 345mm. Length of hind wing 42 mm.
Length of pterostigma 3°75 mm. Breadth of hind wing 13 mm.
Wings hyaline, reticulation black, pterostigma black, mem-
branule brownish grey.
fore wings. 16 antenodals, the last on the right side is dis-
continuous, that on the left continuous. 9 postnodals. Internal
triangle divided into three cells, discoidal triangle crossed by a
single nervule and followed by three rows of cells. Nodal sector
waved at its middle. Two cross nervules in lower basal cells.
Arculus between the level of the first and second antenodal.
Hind wings. 10-11 antenodals, 10-11 postnodals. Discoidal
triangle traversed, followed by two rows of cells. Sectors of
triangle scarcely separate at their origin. Lower basal cell with
a single cross nervule.
Head. WLabrum black, bases of the mandibles bright yellow.
Rhinarium dull yellowish brown, nasus black along its ventral
margin, for the rest yellow. Genz yellow. Frons and tubercle
metallic violet, but frons yellow at the sides. Tubercle truncate
anteriorly, occipital triangle black.
Prothorax brown, posterior lobe with a rounded backwardly
directed projection at the middle of its posterior margin.
Thorax metallic green, marked with dull yellowish brown as
follows :—a fine line along the mid-dorsal carina; a lateral band
74 MR. F. F, LAIDLAW ON THE [Feb. 4,
running from the second and third femora to between the wings.
The whole ventral surface is yellowish brown, and this colour
extends for a short distance along the humeral suture and on to
the sides of the metasternum. Between the wings dorsally are
three yellow spots, one behind the other.
Abdomen black, very slightly dilated at its base. Yellow
lateral spots on segments 1, 2,3. Base of segments 2, 3, 4 with
a fine transverse yellow line; mid-dorsal carina with a scarcely
perceptible yellow line from segments 3 to 7. Longitudinal
yellow marks ventrally on segments 3, 4,5 on either side of the
middle line. Anal appendages black. Transverse carine on
segments 2,3. On segment 2 there is on either side anteriorly a
small tuft of fine black hairs.
Legs black. In the male the second pair of femurs have each
arow of 18 antero-inferior short spines directed towards the knee,
and increasing in size gradually from above downwards. ‘These
are followed after a short gap by three long straight spines which
are also directed a little downwards (see text-fig. 11). The third
pair of femurs have each 26 antero-inferior short subequal spines
all directed towards the knee, save the last six, which are not
inclined. Then follow two longer spines inclined towards the
knee.
Text-fig. 11.
Second femur of Zygonidia malayana.
This species differs from Zygonyx iris chiefly in that the
discoidal triangle of the lower wing is followed by two rows of
cells, not by three; and in the absence of any dorsal markings
on segment 7 of the abdomen, in the coloration of the thorax,
and length of the pterostigma. It is more closely allied to
Zygonidia insignis (Kirby, A. M. N. H. (7) v. p. 540), from
which it is chiefly distinguished by its smaller size, the fewer
reticulations in the postnodal spaces of the hind wings, and in
the details of the spines on the second and third pairs of femurs
of the male. In 4%. insignis the second pair of femurs have
each a row of 18 antero-inferior spines, followed by three much
longer spines. The first 12 are directed towards the knee, the next
six are not inclined, the three long spines are slightly inclined
towards the knee. The third pair of femurs have each some 24
short spines, the first 14 inclined towards the knee, the last 10
scarcely inclined, followed by two longer spines inclined to the
knee.
I caught two specimens of Zygonidia malayana at Kwal
Aring in September. These, like all the other recorded specimens
1902. ] DRAGONFLIES OF THE “ SKEAT EXPEDITION.” (ie)
belonging either to the genus Zygonidia or to Zygonyx, are
both males.
The character of the last antenodal cross nervule seems to be
very variable. In one specimen of Zygonyx iris it is accident-
ally complete (de Selys, C. R. Soc. Ent. Belg. xxxv. p. cexxvii).
In one of the two known specimens of Zygonidia insignis it is
accidentally incomplete on one side (Kirby, A. M. N. H. (7) v.
p- 540). On one side of both specimens described above it is
incomplete, on the other complete.
These two preceding species, together with their allies, pro-
bably constitute a separate section of the Libelluline approxi-
mating somewhat closely to the Corduliine. See Calvert, loc. cit.
ONYCHOTHEMIS TESTACEA, sp. n. (Plate V. fig. 2.)
Length of abdomen 33:5 mm. Length of hind wing 40 mm.
Length of anal app. of g¢ 2°5. Breadth of hind wing 13°5 mm.
Wings hyaline, reticulation black. Pterostigma 4 mm. long.
Fore wings. 15 antenodals, 10-11 postnodals. Internal tri-
angle divided into three cells. Discoidal triangle narrow, with
a single cross vein, followed by three rows of cells: no supra-tri-
angular nervule. Nodal sector waved, a single cross nervule in
the lower basal space; membranule long, grey. Upper sector of
triangle curved. Sectors of arculus stalked.
Hind wings. 9-10 antenodals, 11 postnodals. Discoidal triangle
free, followed by two rows of cells. Lower basal cell with a
single cross nervule. Sectors of triangle originate close together.
Nodal sector waved.
Head. Ventral surfaces yellow, with a triangular black mark
in the centre, its apex directed forwards. Upper lip black, with
a reddish-brown spot on either side. Rhinarium and nasus
reddish brown, with a black mark along the suture between them.
Frons yellow below, steely black above. Frontal tubercle bifid,
steely black. Occipital triangle black.
Prothorax black, with a yellow hinder margin.
Thorax dark metallic green, the mid-dorsal carina and a mid-
dorsal line between the wings yellow. Two small orange-yellow
spots on the humeral sutures on either side, the one above the
other. A thin yellow band runs from immediately behind the
front pair of wings downwards to between the second and third
pairs of legs on either side. An orange line runs along the outer
edge of the metasternum. Ventral surface black.
Abdomen broad, slightly dilated at its base. Segments 4 to 8
strongly triangular in section. Testaceous black with dull yellow
‘markings. Mid-dorsal spots on the middle of segments 1 to 8,
Segments 2 and 3 with a yellow-marked transverse carina; the
yellow mark is discontinuous dorsally in 2, but in 3 runs into
the yellow spot. Ventrally each segment from 3 to 8 has two
large oblong yellowish spots on either side of the middle line.
These spots extend round the lateral keel on to the sides of the
abdomen and on segment 7, 8 reach to within a short distance of
76 MR. F, F. LAIDLAW ON THE [Feb. 4,
the dorsal spots. Segments 9 and 10 black, segment 10 is very
small. Upper appendages black, curved slightly inwards and
at first downwards, but at their distal ends they turn up a little.
About halfway along their ventral sides is a small tooth. Ventral
appendage black, rather broad and flat, bifid at its extremity,
not more than two-thirds the length of the upper pair.
Legs black. On each of the first pair of femurs is a single
Spine immediately before the knee. In the second pair on each
are three long pairs of spines inclined towards the knee and
distant from each other. On each of the third pair of femurs
are 4 pairs of large spines, rapidly increasing distally, followed
after a gap bya single pair. All inclined shghtly towards the
knee. On the first pair of tibias are three pairs of long equi-
distant spines, on the second and third pair of tibias are four
pairs of large spines. (See text-fig. 12.)
Text-fig. 12.
(
Third leg of Onychothemis testacea (X 13).
A single male from Kwala Aring.
This species differs from Onychothemis abnormalis (Brauer,
Verh, Ges. Wien, xviii. p. 170) in having no transverse carina
on the fourth abdominal segment, and in having the claws of the
third pairs of legs without any sign of a tooth. J have not been
able to examine a specimen of O. abnormalis, a Philippine Is.
species, but the present species is evidently closely allied to it.
This genus appears to stand quite remote from other Libel-
lulids, not only in the absence of a tooth on the basal claws, but
in the remarkable armature of the femurs. The two species
of the genus should form an independent subsection of the
Libelluline.
CORDULIINA.
Two members of the subfamily are known to occur in the
Peninsula; these are Macromia westwoodi, Selys, and Idionyx
yolanda, Selys. To these can now be added Macromia ger-
staeckert, recently described by Kriiger from Java (Kriig., Stett.
ent. Zeit. 1899, p. 335), and /dionyx dohrni (loc. cit. p. 326) from
‘Sumatra.
MACROMIA GERSTAECKERI Kriiger.
Macromia gerstaeckeri, Kriiger, Stett. ent. Zeit. 1899, p. 335.
I caught a pair of a species of Macromia, which I refer to this
1902. ] : DRAGONFLIES OF THE ‘‘SKEAT EXPEDITION.” 77
species, at Kwala Aring in September 1899. The male is very
young.
Cie OF
Length of abdomen (without appendages). 36mm. 37:°5mm.,
Jenna Ayah). cotecsct ooutics sandcounnecEanasencee 32 34
LETHE) ROSSI ETONE poco ebboouonocooannocubAer Gnd: ons 2 2
g. Antenodal nervures of fore wing 14, postnodal 7, 3. Ante-
nodal nervures of hind wing 11, postnodal 7, 8, Supra-triangular
nervures of fore wing 3 or 4, of hind wing 2, Nerves of median
space of fore wing 6, of hind wing 4.
The head in this specimen is badly shrivelled, and it is difficult
to make out the characters for the most part; the yellow ‘nasus’
is obvious.
Prothorax dull brown. Thorax iridescent brown, with three
yellow lines on each side. The antehumeral band begins at the
base of the coxe of the first pair of legs and runs about halfway
up the thorax. The middle line commences behind the middle
pair of legs and runs up to behind the front pair of wings, running
right across the back to join that of the other side. The hinder
stripe runs along the hinder outer margin of the abdomen,
starting from behind the last pair of legs. The coxe of the
first pair of legs are yellow, as also the hinder surface of those of
the second and third pairs.
The abdomen is dull brown; segments 1-3 and 7-10 a little
expanded. It has yellow markings as follows :—base of the first
‘segment shading gradually into dull brown at its hinder end,
second segment has a yellow ring covering its anterior half;
third segment has two very small dorsal spots lying very close
together just behind the middle of the segment; seventh seg-
ment with a transverse dorsal band taking up the anterior fifth
of the segment.
Anal appendages subequal, or the inferior a very little larger
than the two upper, about 3 mm. long, as long as the last two
segments. ‘The upper pair have each at about two-thirds of their
length a small tooth on the outer side.
_ 9. Head and thorax as in the male, but the iridescent brown
is replaced in the more adult female by rich metallic green. The
wings have a number of vague brown marks on their outer
halves.
Abdomen. Segments 1-6 metallic black, 7-10 dull black.
Yellow markings as follows :—second segment a long spot on
either side, a transverse line along its anterior margin dorsally,
two small dorsal spots at about the middle of the segment; 3 with
a line on either side along its lateral anterior margin not reaching
the dorsal surface; 7 as in the male, but the yellow mark is
broader, covering about one-fourth of the segment.
As Kriiger has pointed out (loc. cit.), MW. gerstaeckeri differs
_ from its allies in its smaller size; and in the position of the small
second tooth of the superior anal appendages of the male from the
78 MR. F. F. LAIDLAW ON THE [Feb. 4,
other species possessing this character (J/. cincta Rambur, west-
woodi Selys, borneensis, fumata Kriiger), in which the tooth les
at the middle of the length of the appendage.
Ipronyx Dory! Kriiger. (Plate V. fig. 4.)
Idionyx dohrnii, Kriiger, Stett. ent. Zeit. 1899, p. 326.
Q@. Length ofabdomen 26mm. Length of hind wing 26:5 mm. ;
breadth 8-5.
Triangles of all four wings undivided, those of the front wings
with their anterior. borders somewhat broken near their outer
angle. Supra-triangular space crossed by a single nerve in all
four wings. ‘Triangles followed by a single row of cells; in the
hind wings the nerve bordering the first cell of the row ends
against the middle of the outer wall of the triangle, so that the
second cell comes into contact with it (¢f. characters of J. optata
Selys, in Ann. Mus. Gen. (2) x. p. 472). On the margin of the
wings this single row breaks up into 6 cells on the fore, 7 on the
hind wings. The median space in the fore wing is traversed by a
single nerve; hinder wing with two nerves in the median area.
Sectors of arculus with very long stalk, rising in both front and
hind wings almost in the angle made by the arculus and the sub-
median vein. Wings tinged with brownish yellow, which is
darkest at the base.
Head brown, with the upper lip dull yellow. Vertex metallic
blue. Thorax brown-green, with three stripes on each side: the
front one is yellow at the base of the front pair of legs, and fades
into brown as it passes up along the thorax ; the second and third
are yellow. The abdomen is very dark brown, fading into black
at the end; the ventral edge of all the segments except the first
marginal with a fine yellow line, which is broadest on 2-3.
This species differs from all its congeners in having the median
space of the lower wing traversed by two nerves. Itis also a trifle
smaller than the other species.
One female from Kwala Aring taken in September 1899.
I have given the characters of this and the preceding species
rather fully because they show some points of variance with the
types described by Kriiger, and in the case of the first between
the male and female.
Family ASSCHNIDA.
Subfamily AU.scHNIN4.
ANAX GUTTATUS (Burm.).
Anaz guttatus, Kirby, Cat. Odonata, p. 84.
One specimen from Kwala Aring. Widely distributed in the
East Indies.
AMPHLESCHNA AMPLA (Ramb.).
Amphieschna ampla, Kirby, Cat. Odonata, p. 93 ; Karsch, Ent.
Nachr. xvii. (1891) no. 18, p. 10.
1902.] DRAGONFLIES OF THE “ SKEAT EXPEDITION.” 79
Two males of this species were taken at the foot of Gunong
Inas. The larger specimen had its abdomen 64 mm. long, its
hind wings each 62mm. The upper pair of abdominal appendages
measure 6°55 mm. This is a very handsome creature, the rich
green and black thorax contrasting with the black yellow-ringed
abdomen. Previously recorded from Java and Amboyna.
TETRACANTHAGYNA PLAGIATA (Waterh.).
Tetracanthagyna plagiata, Kirby, Cat. Odonata, p. 94.
Gynacantha plagiata, Karsch, Ent. Nachr. xvii. (1891) no. 18,
1Do .
I caught a female of this splendid species at Kwala Aring, in
thick forest. It agrees very closely with the type specimen
figured by Waterhouse, from Borneo (Proc. Ent. Soc. Lond. 1877,
p. x; Trans. Ent. Soc. Lond. 1878, p. 119, fig. 4), but is somewhat
smaller. The length of the hind wing is 69 mm., breadth 22 mm. ;
length of abdomen 56 mm.; breadth of head 15 mm. Recorded
from Borneo and Sumatra.
My specimen hasa small supernumerary spine on the right side
in addition to the four normal spines on the end of the abdomen.
GYNACANTHA ROSENBERGII Brauer.
Acanthagyna rosenbergti, Kirby, Cat. Odonata, p. 95.
Gynacantha rosenbergii, Karsch, Ent. Nachr. xvi. (1891)
no. 18, p. 9; Kriiger, Stett. ent. Zeit, 1898, p. 278.
A single male and two females from Kwala Aring. Two other
females probably belong to another species, but I have not been
able to compare them with a series. They are from the same
locality as the rest.
*JAGORIA P@CILOPTERA Karsch.
Jagoria peciloptera, Kirby, Cat. Odonata, p. 91; Kriiger, Stett.
ent. Zeit. 1898, p. 329.
Recorded by Kriiger (loc. cit.) from Singapore.
Subfamily GoMPHINA.
Apparently only three species of Gomphine Dragonflies have
hitherto been recorded from the Peninsula. These are :—
Legion Gompuus.
Microgomphus chelifer Selys (Mt. Ophir, Sumatra).
Macrogomphus thoraciews MecLach. (Perak).
Legion LINDENIA.
Ictinus melanops Selys (Malacca).
Our collection contains five specimens referable to the following
species :—
Legion GomPuus.
Gomphus consobrinus ¢ 2, sp. n.
80 MR. F. F, LAIDLAW ON THE [ Feb. 4,
Onychogomphus geometricus, var. nigrescens n. (Kwala Aring,
Kelantan), @. -
Legion GoMPHOIDES.
Steboldius grandis ( ¢, Gunong Inas, Perak).
Legion LixpENTA.
Gomphidia perakensis, sp. n. (¢, Gunong Inas, Perak).
Legion GomPuvs.
GOMPHUS CONSOBRINUS sp. n. (Plate V. fig. 5.)
Length of abdomen 31 mm. Length of hind wing 26 mm.
Length of pterostigma 2 mm.
Fore wing. No basal subcostal nervule. Sectors of the arculus
distinct at their origin, then converging, meeting for a short
distance, then diverging’. 11 antenodals, 10 postnodals. Ptero-
stigma brownish black, thick. ‘Triangles free,
Hind wing. 10 antenodals, 10 postnodals.
Reticulation black, membranule almost entirely absent.
General colour dull bronze-brown, lower surfaces of head and
thorax greenish yellow. Dorsal surface of thorax dark brown,
with two lighter submedian somewhat oblique bands diverging
from each other from above downwards, each joining a lighter
transverse antehumeral mark so as to make a 7 on either side.
Sides of thorax lighter bronze.
Abdomen almost black, first segment greenish yellow; sides of
second segment, a very fine dorsal longitudinal line on the second
segment, and the auricles yellow. Segments 1, 2 somewhat
dilated, 3 to 7 thin cylindrical, 8 to 10 dilated and progressively
shorter.
Appendages black, upper pair of about the same length as the
tenth segment, widely diverging from each other, turned up and
pointed at their ends, each with a small tooth on the margin at
about half its length. Lower pair diverging, rather shorter than
upper pair, each terminating in a laterally directed point.
A male and a female were taken at Kwala Aring. The female
had only very recently escaped from the larva and is too much
withered to describe.
There seems no doubt that this is a true Gomphus belonging
to Type A of Selys (Mon. Gomph. p. 376). The occurrence of a
member of this group in such a locality is remarkable.
ONYCHOGOMPHUS GEOMETRICUS NIGRESCENS, var. n.
One 9 from Kwala Aring, Kelantan.
Closely allied to O. geometricus de Haan.
Head, prothorax, and thorax as in O. geometricus.
Posterior surface of first pair of femurs yellow.
Abdomen black, marked with yellow as follows :—a dorsal band
on the first and second segments, decreasing in size from before
backwards, the sides of these segments are yellow. The auricles
1 This feature is not shown in the figure.
1902. ] DRAGONFLIES OF THE “‘SKEAT EXPEDITION.” 81
are scarcely visible. Segments 3-7 have an incomplete basal
orange ring, occupying in segment 3 about one-fourth of the
segment, in 4-6 not more than one-sixth, and in 7 not more than
two-fifths of the segment. Remaining three segments black,
decreasing successively in size; anal appendages pointed, larger
than the tenth segment, yellow.
Differs from the typical race in the smaller extent of the yellow
marks on the abdomen. The yellow lozenge-shaped dorsal spot
on segment 3 is present, that on 4 is scarcely visible. The great
length of the first two cells between the sectors of the arculus is
remarkable. The cross nervule enclosing the third cell is beyond
the level of origin of the principal sector.
Legion LINDENIA.
GOMPHIDIA PERAKENSIS, sp, n. (Plate VI. figs. 1, 2.
Rotalolemothae: jac: sy seeeeere tec cess 78 mm.
Length of abdomen (without ap-
pendages)|. stra ceeaes ese n ts wttleceis A
Breacdiblav ol Weadvaenaaeeee se ch ete ese PUBS) ss
Length of hind wing.................. 4,
ute eg OLE SWUNG. waits de sele ne eons Bl ce
Rs A ePUCEOSUICTMAN 5.2. 8ee ace = Siar.
Breadthvot bund! wang -2-...--- irae
Length of upper anal appendage... fe:
Colour black with saffron-yellow markings; wings hyaline,
reticulation black.
Fore wing. 22-23 antenodals, 17-18 postnoda’s. Internal
triangle divided into three cells; discoidal triangle into four.
Membranule small, dark brown. Pterostigma long, black.
Hind wing. 15-16 antenodals, 20-21 postnodals. Internal
triangle divided into three cells ; discoidal triangle into four,
Head. Labium, mandibles, and labrum black; gena black, with
a dull yellow spot. Rhinarium saffron-yellow ; nasus black, with
a saffron-yellow spot on either side. Anter:or surface of frons
black, horizontal surface yellow. Vertex and occiput black,
The vertex has two conical projections, one on either side as in
G. T-nigrum, but not so large relatively. Prothorax black.
Thorax black, with the following saffron-yellow marks :—In front
dorsally a semicircle broken by the mid-dorsal carina. From the
outer ends of the semicircle, but separated widely from them, a
short broad band runs on either side to the ante-alar sinus,
inclining towards the middle lines. Behind these on either side
is a small spot, just below the sinus and immediately in front of
the first pair of wings. Laterally are two large bands widely
separated. The first pair run obliquely forward from below the
first pair of wings; the second pair are rather smaller, and run
obliquely forward from below the second pair of wings: neither
pair reaches the ventral surface.
Dorsally between the wings there are brown marks and a
Proce. Zoou. Soc.—1902, Vou. I. No. VI. 6
82 MR. F, F. LAIDLAW ON THE [Feb. 4,
citron-yellow spot between the first pair of wings. Ventral
surfaces and legs rich black. The legs are robust, rather short ;
hindermost pair of femurs with stout spines.
Abdomen somewhat dilated at its base, then segments 3 to 6
long and cylindrical, segments 7 to 9 compressed laterally, 10 very
short. Black, with yellowish marks as follows :—on segment 2 a
lateral spot, the auricles, and a very small dorsal lozenge-shaped
mark, Segment 3 has a trace of a yellow spot on either side and
a very fine mid-dorsal line. The basal third of segment 7 with a
large dorsal mark. Anal appendages black, resembling those of
G. T-nigrum very closely.
Coloration generally remarkably similar to that of a specimen
marked Macrogomphus quadratus in the British Museum.
A single male from forests at the foot of Gunong Inas. This
fine species differs from other members of the genus Gomphidia
in having its wings of equal length with the abdomen. It
appears, moreover, to be the largest known member of the
legion Lindenia. The internal nervule of the pterostigma can
hardly be said to be prolonged, but this character is scarcely of
sufficient importance to justify the removal of this species from
the genus Gomphidia. The form of the anal appendages and the
absence of any leaf-like dilatation of the sides of segments 7 or
8, as well as the shape of the vertex, indicate that it must be
referred to this genus, of which it may form a new section.
Legion GoMPHOIDES.
SIEBOLDIUS GRANDIS Kriiger. (Plate VI. figs. 3, 3a, 4.)
Sieboldius grandis, Kriiger, Stett. ent. Zeit. 1898, p. 311.
I refer to Kriiger’s species a fine male, which agrees in size,
wing-characters, and coloration fairly closely with the females
described by him from Sumatra.
Motalelenetnet, Hywel ser oes 83 mm.
Length of abdomen (without ap-
PEMCAGER) UM... acme ends oo ear:
Length of appendages (upper pair). Qs,
Hore wine lene th eereyscees. aarti Does.
x‘ breachths nis, ates, Gee aes
iEindawanies Teme theres seeceeectae cater Daas
-Y bveaditth, (225. 325. Seas LS tes
Pieros hima MI wKI5 eo IGE ce Pe DAD
Femur of last pair of legs............ 20° 3
Tibia a gy 0 ELT EARN Ts wy
iBréadthof head 2s. ajecs-est tecacee 11
Antenodal cells of fore wing 22—23, postnodal 18-19; of hind
wing 16-17 and 17 respectively. Pterostigma lies over six cells.
Basal subcostal nerve present in all wings. Triangles of all four
wings with one cross nervure. The middle thirds of all the wings
1902.] DRAGONFLIES OF THE ‘‘ SKEAT EXPEDITION.” 83
have, when looked at obliquely, a very faint whitish “smoky ”
appearance.
Head small, black save for a yellow transverse band on the
‘frons’ before the eyes, stopping abruptly at its anterior edge.
Eyes distant. Occiput with the two convexities at its hinder
margin more pronounced than in S. japonicus.
Prothorax black; at its hinder margin a transverse yellow
band tapering laterally. In front of this 1s a yellow spot.
The colouring of the thorax is just as described by Kriiger for
the female. On the upperside a yellow band runs from the front
margin up to the yellow marking in between the wings; this
band is twiceas broad in front end as it is at its hinder end. On
either side are two fairly broad yellow oblique bands,
The abdomen is ringed with black and yellow; the first segment
is yellow with a black mark on either side. These black marks
are continued on to the second segment, in which they run a
little obliquely up to the dorsal surface, meeting at the hinder
end of the segment; a fine black ring runs round its posterior
margin. Thissegment is black below, and the auricles are tipped
with black continuous with the lateral stripes. The yellow ring
of segments 3-8 occupies the following portion of each segment :
the front two-fifths of 3, the front one-third of 4, 5, one-fourth
of 6, 7, two-fifths of 8; 9 and 10 are entirely black. There isa
very fine mid-dorsal black line in 3; this is present, increasing
in breadth as one passes back from 4-7, but absent in 8. ‘The
appendages of the tenth segment agree very closely w.th those
figured by de Selys for S. japonicus (Selys, Mon. Gomph. pl. xu.
fig. 3b). The upper pair are rather shorter than the tenth
segment, slightly ciliated except at their end, which is sharply
pointed and curved upwards ; they carry two teeth on their lower
side, one at about a third of their length rather blunt, directed
downwards and a little outwards, the other at the end of the
second third, sharper and curved backwards. The lower pair
about half the length of the upper pair, thick and blunt.
The femurs of each leg have a number of short prickly spines
on their outer lateral face; the hinder pair have a few delicate
hairs on their upper surface. All have short tooth-like spines in
regular rows along their lower sides.
A single specimen (¢) was caught at the foot of Gunong Inas
(about 1000 feet above sea-level) near a small jungle-pool, in
January 1900.
Family CALOPTERYGID.
Subfamily CALOPTERYGIN &.
The following is a list of the Calopterygines mentioned in
Kirby’s Catalogue or elsewhere as known to oceur in the Malay
Peninsula up to 1890:
Neurobasis chinensis Linn.
Vestalis amena Hagen.
6*
84 MR. F. F. LAIDLAW ON THE [Feb. 4,
EHuphea impar Selys.
5 ochracea Selys.
Dysphea limbata Selys.
Devadetta argyroides Selys.
Rhinocypha fenestrella Selys.
A biforata Selys.
i petiolata Selys.
Micromerus aurantiacus Selys.
3 stigmatizans Selys.
a hyalinus Selys.
Since that date Dohrn has added the following to the list :—
Micromerus lineatus Selys.
s signatus Kriiger.
Our collection contains examples of the species enumerated
below :—
* cho modesta 2 , sp. n.
*Climacobasis lugens 3, sp. N.
Neurobasis chinensis 3 2.
Vestalis amena 3 Q.
Euphea impar 3.
» ochracea 3 Q.
Dyspheea limbata 3.
Rhinocypha fenestrella $ 2.
se biforata 3 Q.
A inas ¢ 2, sp. n.
. 4 karschi 3.
* Micromerus affinis 3 9, sp. n.
[Species marked * are new to the Peninsula. |
*
Legion CALOPTERYX.
EcnHo MoDESTA, sp. n. (Plate V. fig. 6.)
1 9, Kwala Aring.
Abdomen, length 41 mm. Hind wing, length 37:5 mm.
Fore wing with 33-36 antenodals, circ. 45 postnodals: hind wing,
32 antenodals and cire. 40 postnodals. Basal area with 9-10 cross
nerves in fore wing, 8-9 in hind wing. Quadrilateral with 9-10
cross nerves in fore wing, 7 in hind wing.
Head. Mouth-parts black; antenne black, second joint long
and thick, third joint longer but much thinner. Rhinarium
black, nasus bright metallic green, rest of the head very dark
bronze-green.
Prothorax dark bronze-green. Thorax the same colour, rather
brighter at the sides; underparts brown; legs of the same
colour, but the femurs have some irregular black marks on their
upper sides ; hairs very long and numerous.
Wings hyaline, with a faint brownish tinge at their outer
extremities. Pterostigma rather longer than broad, pale brown,
1902. ] DRAGONFLIES OF THE “‘ SKEAT EXPEDITION.” 85
lying over 4-6 cells. Behind the pterostigma are first two rows
of cells, then, after about five cells, only one row.
Abdomen dark brown, with a green iridescence in some lights
on the first three and last three segments. Tenth segment very
short, not half as long as the ninth, ninth longer than the eighth.
Appendages shorter than the tenth, black, conical, and sharply
pointed.
This species differs from H. uniformis Selys (? = £. tricolor
Kriiger) in having a smaller number of postnodal cells (it pos-
sesses 45 as against 60-65), in its rather smaller size, and in
the colouring of the wings, which are described by Kriiger as
being in the female yellow all over, especially at the base and
anterior margin, whereas in /. modesta the base of the wings is
perfectly transparent (Kriiger, Stett. ent. Zeit. 1893, p. 72; Selys,
Bull. Ac. Belg. (2) xlvii. p. 357, id. Ann. Soc. Ent. Belg.).
There is a female in the British Museum from Mr. Ridley,
collected in Penang, belonging to this species. It has 37 ante-
nodals and 48 postnodals in the fore wings. Its abdomen is of a
dull dark red-brown colour.
CLIMACOBASIS, gen. nov.
Basal area of wings reticulated. Pterostigma long; quadri-
lateral long, rectangular; arculus bent, sectors starting at the
same point just below its middle. Principal and subnodal sectors
rise at about the same level from the reticulum, between the
upper sector of the arculus and the median nerve.
The nervule closing the lower basal cell runs from the lower
sector of the arculus straight down to the lower extremity of the
lower basal cell.
CLIMACOBASIS LUGENS, sp. n. (Plate VI. fig. 5.)
(Last three segments of the abdomen missing.)
Length of abdomen (segments 1-7) 42 mm. Length of hind
wing 47 mm. Breadth of hind wing 10 mm.
Fore wing with 37 antenodals, 45 postnodal nerves. Ptero-
stigma covering 8-9 cells, about 2°5 mm. in length, very black.
Basal area with 8 cross nerves.
Hind wing. 34 antenodals, 37 postnodals. Pterostigma as in
fore wing, basal area with 8 cross nerves.
Head. Lower lip, base of the mandibles, and upper lip black.
Between the eyes, running forward as far as the epistome, is a
remarkable square milky-white patch of considerable size, taking
up in fact the greater part of the vertex. Along its hinder margin
it is notched in the middle by the anterior ocellus, which is sur-
rounded by a very small black ring which is continuous with a
rectangular black patch, in which lie the two posterior ocelli; the
rest of the head is of a very dark bronze-green colour.
Prothorax dark green, almost black.
Thorax. Dark metallic green above, with all the sutures and
86 MR. F. F. LAIDLAW ON THE ’ [ Feb. 4,
the interalar space black; under surface and legs sooty black
with long hairs. Wings hyaline. Legs brownish black, with
very long hairs.
Abdomen (first 7 segments only) dull brownish black.
There can be, I think, no doubt that this species has as its
nearest known ally Archinewra. I believe, however, the differences
between them are of generic rank, the chief of these being the
much smaller number of accessory nervures running to the hinder
margin of the wings, the mode of origin of the principal and sub-
nodal sectors, and the character of the nerve running to the lower
basal cell. ;
The only specimen taken has unfortunately been rather badly
knocked about and has lost the last segments of the abdomen. It
was caught in September in jungle at Kwala Aring. Its habits
were s-milar to those of Vestalis amena, for which at first I mis-
took it.
Foerster, in discussing the affinities of the genus Matronoides,
has proposed the following arrangement of the genera belonging
to the legion Calopteryx (Foerster, Ann. Soc. Ent. Belg. p. 66,
1899) :—
(Syz phis.
Calopteryx.
“| Phaon.
} Vestalis.
(No pterostigma ......
Basilar space free
| ( Umma.
, } Sapho.
. . r re
( Pterostigma present 4 Mnais.
(_Psolodesmus.
; Matrona.
[ No pterostigma ......... Neurobasis. ~ Matronoides.
Basilar space with cross Neurobasis.
nervules.
( Echo.
U Archineura.
LA pterostigma present.
If this grouping be accepted, and it is very convenient, the last
division may now stand as follows :—
(Short rhomboidal pterostigma... Hecho.
Pterostigma present. Pterostigma at least three Archineura.
{times as long as broad. Climacobasis.
NEUROBASIS CHINENSIS (Linn.).
Weurobasis chinensis, Kirby, Cat. Odonata, p. 102; Selys,
Odon. de Sumatra, Ann. Mus. Genova (2) vii. p. 189; Selys, Odon.
de Birmanie, loc. cit. (2) x. 1890-1, p. 487; Selys, Vewrobasis
chinensis et ses races locales, Ann. Soc. Ent. Belg. 1896; Kayrsch,
Ent. Nachr. xvi. no. 16, p. 243.
Five males, three females, from the Aring River in Kelantan.
This species appears to travel further down the rivers than any
other Calopterygine, at least so far as my observations went. It
is very widely spread in Tropical Asia.
1902. ] DRAGONFLIES OF THE “‘ SKEAT EXPEDITION.” 87
VESTALIS AM@NA Hagen.
Vestalis amena, Kirby, Cat. Odonata, p. 103; Karsch, Ent.
Cotes xvii. 1891, no. 16, p. 242; Kriiger, Stett. ent. Zeit. 1898,
p. 70.
Several males and females from Kwala Aring and from the
foot of Gunong Inas.
This species occurs also in Borneo, Java, and Sumatra.
Note.—Two females from Kwala Aring differ rather markedly
from the rest of our specimens. The general colour of the body
is dark bronze-green rather than emerald-green of the other
specimens. Further, the wings have a distinct brownish tinge.
In respect to the markings on the head, the yellow is brighter
than in the other specimen. One male shows a tendency to have
the wings tinged and is also of a more bronze-green shade than
the other males. These three specimens are perhaps much more
adult than the others.
Legion Evry”.
Genus EKupH#A Ramb.
ELuphea Ramb. Ins. Névr. p. 228 (1842); Selys, Syn. Cat., Bull.
Ac. Belg. 1853, p. 50; id. Mon. Cal., l.c. 1854, p. 167.
Pseudophea Kirby, Cat. Odonata, p. 109.
Huphea Selys, Ann. Soc. Ent. Belg. p. 338 (1891).
EvupH#A IMpar Selys.
Pseudophea impar, Kirby, Cat. Odonata, p. 109.
Four males from the Aring River above Kwala Aring.
This species differs greatly from the following, not merely in
the colouring of the wing but also in the wings being much
broader proportionately and with very rounded tips.
Eupu@A ocHRACEA Selys.
Pseudophea ochracea, Kirby, Cat. Odonata, p. 109.
Huphea ochracea, Selys, Ann. Mus. Genova, (2) x. p. 489.
Four dg, one 2: two males from the Aring River in Kelantan,
the other three individuals from the Selama River at the foot of
Gunong Inas.
The male is a very beautiful insect, and when alive his wings
seem to be almost crimson in colour as he hovers over the surface
of the stream. The wings of the female specimen have hardly a
trace of yellow tinge (cf. Selys, loc. cit.), and the pterostigma is
brown, those of the wings of the male being rich velvety black.
The rich red markings of the thorax of the male are dull brown
in the female, and the whole body is duller,
Length of abdomen without Ou: De
appendages ............... 36mm, 30 mm.
Length of hind wing......... 20. DOM Oe.
i Fe MOSHE) WU? Boscadnte: oie, 307). 4,
Known also from Burmah and Borneo.
88 MR. F, F, LAIDLAW ON THE [Feb. 4,
DysPHaA LIMBATA Selys.
Dysphea limbata, Kirby, Cat. Odonata, p. 110.
Seven males from the Aring River some way above Kwala
Aring. Known also from Borneo.
This species is regarded by Selys as a local race of D. dimidiata
Selys, described from Borneo. In all my specimens the black
mark at the base of the fore wing extends just beyond the level
of the nodus and its margin is straight, at right angles to the
anterior margin of the wing. On the hinder wing the black
basal mark reaches halfway between the nodus and pterostigma
and its outer margin slopes inwards a little.
Selys has remarked on the scarcity of the females of this group
(Bull. Ac. Belg. (2) xxxv. p. 487). From my own experience I
am sure that this scarcity in collections is not due to their being
overlooked by collectors. JI can safely say that I never saw a
female of this species or of Huphea impar, whilst the males were
at times abundant.
Legion LiBELLAGO.
RHINOCYPHA FENESTRELLA Ramb.
Rhinocypha fenestrella, Kirby, Cat. Odonata, p. 113; Selys,
Ann. Mus. Genov. (2) p. 491 (1891).
This species, which is closely allied to R. quadrimaculata of
India and &. spuria of the Khasia Hills, ranges from Burmah as
far south as Penang. It is fairly common on the Kelantan River,
and Mr. Evans took some specimens in Patalung. There are two
specimens in the British Museum from Province Wellesley, taken
by Mr. Ridley.
RHINOCYPHA BIFORATA Selys.
Rhinocypha biforata, Kirby, Cat. Odonata, p. 113.
This species occurs rather more abundantly than the last on
the Kelantan River. I also found it fairly common near the
foot of Gunong Inas.
RHINOCYPHA INAS, sp. n. (Plate VI. fig. 6.)
Length of abdomen g 19mm. Length of hind wing ¢ 23 mm.
Length of abdomen 9 18mm. Length of hind wing 9 24 mm.
3. Black. Head with five yellow spots, two in front on either
side of the ocelli, three in a transverse row behind these, the
median spot transversely elongated.
Prothorax with two small anterior and two larger lateral blue
marks. Posterior lobe orange with black margin.
Thorax with a short blue mesothoracic triangle rose-colour,
on either side of this lies a blue triangular humeral mark not
extending higher than the apex of the mesothoracic triangle.
Sides of thorax blue with two black marks on either side, the
anterior runn'ng from below the first pair of wings does not
extend all the way down to the legs. The second stripe, which is
1902. DRAGONFLIES OF THE “‘ SKEAT EXPEDITION.” 89
broader above, runs from below the hinder pair of wings to behind
the third pair of legs. The blue sides are margined postero-
ventrally with a black line. Lower surface black, with two broad
blue marks behind the legs.
Abdomen black. Segments 1-9 with triangular blue spots on
either side, their bases resting on the hinder margin of each
segment. Those in 2-3 extend the whole length of their seg-
ments, that in 4 for half the length of that segment, the rest are
small.
Anal appendages black. Legs black. The two hinder pairs of
femurs and tibias are white on their inner surface.
Wings tinged with yellow, blackish in the costal area from the
fifth postnodal cross nerve; from about three-fifths the distance
between the nodus and the black pterostigma the apical portion
of the fore wing is purplish brown to the tip, except along its lower
margin, where it is dusky grey shot with iridescent green. The
purple mark commences suddenly and its inner margin slopes
outwards from in front.
The outer half of the lower wing is also marked with brownish
purple, the extreme apex and the posterior margin excepted, these
are greyish brown. The inner border of the purple mark is
straight, and the mark is crossed by two rows of iridescent
hyaline spots. The inner row consists of three spots. The upper
of these, consisting of one row of cells, lies above the nodal sector ;
the second, consisting first of one and later of two rows, lies between
the subnodal and median sector; and the third, above the upper
sector of the triangle, consists of a single row of cells. The upper
spot is nearest to the base of the wings, the lowest is furthest
from the base.
The first row lies at a level of about half of the distance
between the nodus and pterostigma. The second row consists
also of three spots. The upper is the largest and its distal end
just overlaps the pterostigma. It consists of two, followed by
three or four rows of cells, and is placed in series with the upper
spot of the first row. The lowest spot of the outer series is placed
serially with the middle spot of the inner series, whilst the middle
spot of the outer series, consisting of a single row of cells, lies
between the two others.
There is also a hyaline spot just at the middle of the hind wing
consisting of a single row of cells, this impinges on the inner
margin of the brown spot.
2. Head as in the male, with the following additional marks
yellow :—four spots on the dorsal surface of the ‘snout’ and the
second joint of the antenne, the upper half of the epistome.
The gene are marked with greenish blue in the male and yellow
in the female. Prothorax black, with lateral yellow and a fine
mid-dorsal yellow spot.
Thorax black, mesothoracic carina orange; a fine orange line
running to the base of the first pair of wings between the carina
and the humeral suture on either side, this latter is also yellow;
90 MR. F. F. LAIDLAW ON THE [Feb. 4,
under surface yellow, the yellow extends for a short way on to
the sides. Abdomen black, mid-dorsal carina orange, a yellow
spot on either side of first segment. Segments 2-4 have on
either side a yellow line followed by a yellow dot. Wings
hyaline.
Antenodals 11-14.
This species is closely allied to 2. perforata, but differs from
that species and its other allies in the greater extent of the
purple mark on the fore wings of the male. The marks of
the hind wing resemble most closely those of 2. whiteheadi Kirby.
Seven males, four females, Gunong Inas.
RuINOCYPHA KARSCHI Kriiger, Stett. ent. Zeit. 1898, p. 33.
Three males from the Aring River near Kwala Aring.
Abdomen, length without appendages ...... 14°15 mm.
METiinn el WAM sr aac. ate ee et one neo cee meena 20 E.
A single row of postcostal cells. Fore wings yellowish hyaline,
hind wings with a blackish-brown mark covering their outer
extremities starting about halfway between the nodus and ptero-
stigma, its inner margin convex. No vitreous spots on the wings.
The abdomen has on its dorsal side a brick-red spot on segments
2-6; that on segment 2 is small and oval, from 3-6 the spots
are large and rectangular, the sides of the rectangle are longer
than the ends. On segment 7 are two long red lines divided by
a fine mid-dorsal black lime, and on segment 8 two very small red
spots similarly divided. In one case the spot on segment 6 is
also divided by a black line. The sides of segments 1-8 have
each a yellow comma-shaped mark.
Kriiger points out that this species belongs to a group inter-
mediate in character between &. heterostigma and RK. tincta of
de Selys. Described by Kriiger from specimens from Sumatra.
MICROMERUS AFFINIS, sp. n. (Plate VI. fig. 7.)
Two males, one female, Kwala Aring.
3. Head black, rhinarium dark metallic blue. A small reddish-
yellow spot on either side of the ocelli, behind these three others
of the same colour, viz. a transversely elongated median spot on
the top of the occiput and two lateral spots.
Prothorax black, with two small lateral spots, two anterior
dorsal spots, and a single posterior dorsal spot of the same yellow
colour as the head-spots. There is also a fine yellow line running
along the dorsal posterior margin, ending laterally in a spot of the
same colour just above the base of the first pair of legs.
Thorax black, a small antehumeral stripe on either side not
reaching to the top of the mesothorax, and a fine line on the
upper half of the humeral suture also yellow. At the sides are
two large oblique yellow bars; the anterior of these is divided
into two halves by a black mark projecting into it from its hinder
1902. ] DRAGONFLIES OF THE ‘‘ SKEAT EXPEDITION.” 9]
margin. In the upper half is a large black spot, and at the bottom
of the lower half is a smaller spot also black.
Wings hyaline. Outer two-sevenths of fore wings (which are
without pterostigma) opaque dark brown. Five antenodals (six
in one case). Lower wings with a slight brownish opacity at the
margin. Postnodals 11—13.
Length of hind wing 17 mm.
Abdomen black, with the following yellow marks:—On seg-
ments 2-6 a dorsal spot divided longitudinally into two by the
mid-dorsal black carina. On segments 4-5 these marks have an
anterior lateral prolongation, giving them the appearance of two
figure 7’s lying back to back; in segment 6 the lower limb of
the 7, so to speak, has disappeared, leaving merely two anterior
marginal lines. A large lateral spot on segment 1. On either
side of segments 2—3 are two spots in the form of a !, the ‘dash’
being anterior. On 4-5 only the ‘dash’ is present and is very
small on 5. Anal appendages black, upper pair two-thirds length
of the 9th segment.
@. Head as in the male, with black less velvety, yellow marks
lighter, and the following. additional yellow markings :—Basal
parts of lower lip and of labrum, epistome with a yellow spot on
either side. Frons with four spots arranged in the form of a
square, the anterior pair larger than the posterior. Genal region
yellow.
Prothorax and thorax as in the male, but the thorax has a
median dorsal yellow stripe. Abdomen dull black, with a fine
mid-dorsal yellowish line on each segment, not continuous at the
margins from segments 2-9. In 9 it occupies only the posterior
half of the segment. Segments 1—9 each with a conspicuous
lateral yellow mark running nearly the whole length of the seg-
ment, narrowest at the middle except in 9, where it is reduced
to a spot at the hind end of the segment, which is larger than
the eighth, as large as the seventh. Tenth very small. Length
of abdomen 13 mm.
Wings hyaline, pterostigmata pale brown. Length of hind
wing 19 mm.
Differs from J/. semiopacus in having the apex of the hinder
wing opaque, in the possession of markings on the head, and in
the spots on segment 6 of the abdomen. The brown mark of the
front wings is also rather less extensive, 5; mm.; 6 in JZ, semi-
opacus. From MM. martine Karsch it differs in having only
three yellow spots at the back of the head, in the markings at the
sides of the thorax, and in haying the dorsal abdominal markings
broader in front; also in the rather smaller number of ante-
nodal nerves on the fore wing.
92 ON THE DRAGONFLIES OF THE “‘SKEAT EXPEDITION.” [Feb. 4,
Description of a new Species of the Genus Lestes.
LESTES RIDLEYI, sp. 0.
3. Length of abdomen (without appendages) 48 mm., of hind
wing 31 mm., pterostigma 2°75 to 3 mm.
General colour dull bronze-green. Wangs hyaline, iridescent,
slightly tinged with brown at the tip. Two supplementary
sectors between the subnodal and median sector, 18-19 post-
nodals. The nodal sector begins in the seventh cell after the
nodus in the fore wings, and in the sixth in the hind wings.
Reticulation and pterostigma black.
Head. Lower lip dull yellowish brown, upper lip brown, rest of
the upper part of the head bronze-coloured.
Prothorax dull brown, with a small bronze transverse mark
along its posterior margin, which is not indented.
Thorax. Upper surface dark bronze-green, with an obscure
paler line following the humeral suture. Sides and lower surface
yellowish brown.
Abdomen. First segment yellowish brown. The segments 2-7
have a roughened dorsal surface, which is brown-green with a
fine yellowish-green basal ring; ventral surface bluish green, the
bronze extends laterally ; segments 8-10 smooth, bluish, pruinose.
Anal appendages lost, but, if I remember rightly, these were also
of a bluish colour.
The male was taken in the same locality as the specimen of
Pericnemis, at the foot of Gunong Inas. There is stated to be
a female belonging to the same species, which I have not ex-
amined, in the British Museum collection, taken by Mr. Ridley
in Singapore.
L. ridleyi is closely allied to L. orientalis Hagen, from Ceylon,
and Z. udeana Kriiger, from Sumatra. It is sufficiently distin-
guished from both by its size, being intermediate in this respect.
EXPLANATION OF THE PLATES.
PuatTE V.
Fig. 1. Neurocena ida 9, p. 72.
2. Onychothemis testacea 3, p. 75.
3. Tetrathemis pulchra a p. 71.
4. Idionyx dohrni 9, p. 7
5. Gomphus consobrinus = p. 80.
6. Echo modesta &, p. 84.
Puate VI.
Fig. 1. Gomphidia perakensis 3, p. 81.
2. End of abdomen of ditto.
3. Sieboldius grandis g, p. 82.
3a. End of abdomen of ditto.
4, Side view of thorax of ditto (Xx 14).
. Climacobasis lugens, ae cme: p. 86.
. Rhinocypha inas $,p
<7. Micromerus affinis 7 io 14), p. 90.
Ee
1902.] ON ORTHOPTERA FROM BRITISH E. AFRICA AND UGANDA. 93
3. List of a small Collection of Orthopterous Insects formed
by Sir Harry Johnston in British Hast Africa and
Uganda in 1899 and 1900, with Descriptions of Five
new Species. By W.F. Kirsy, F.L.S., F.E.S., Assistant
in the Zoological Department, British Museum (Natural
History), South Kensington.
{Received November 28, 1901.]
The total number of species of Orthopterous Insects repre-
sented in the collection is 27, of which 23 are enumerated in
the present paper, four species, probably new, remaining over for
future consideration.
BuATTID&4.
BLATTIN2.
DEROPELTIS.
Deropeltis Burm. Handb. Ent. ii. p. 486 (1838).
1. DEROPELTIS MELANOPHILA.
Ischnoptera melanophila Walk. Cat. Blatt., Suppl. p. 146 (1869).
One male, from Baringo, 4000 feet, Dec. 20, 1899.
This species was described by Walker from Zanzibar. There
are also specimens in the Natural History Museum from
Samburu, British East Africa, from Mr. C.8. Betton’s collection,
and from Mombasa and Madagascar. This species differs from
D. erythrocephala Fabr. by the black head, with only a streak
within the antenne, and the lower mouth-parts red.
POLYPHAGINE.
POLYPHAGA.
Polyphaga Brullé, Hist. Nat. Ins. ix. p. 57 (1835).
Heterogamia Burm. Handb. Ent. 11. p. 488 (1838).
2. PoLYPHAGA AGYPTIACA.
Blatta egyptiaca Linn. Syst. Nat. (ed. x.) p. 424. n. 2 (1758).
One female specimen, Baringo, 4000 feet, Dec. 20, 1899.
Widely distributed in Africa and Southern Europe.
MANTID &.
MANTIN#.
TENODERA.
Tenodera Burm. Handb. Ent. 11. p. 534 (1838).
3. TENODERA CAPITATA.
Tenodera capitata Sauss. Mitth. Schweiz. ent. Ges. ili. p. 69
(1869) ; Mém. Soc. Geneve, xxi. p. 293 (1871).
94 MR. W. F. KIRBY ON ORTHOPTERA FROM [ Feb. 4,
Mount Ruwenzori; a large specimen measuring 53 inches in
expanse.
This species inhabits Hast and Central Africa and the Congo
district.
VATINE.
Popa.
Popa Stal, Gifv. Vet.-Akad. Forh. xiii. p. 169 (1856).
4, Popa UNDATA.
Mantis undata Fabr. Ent. Syst. ii. p. 19 (1798).
Mount Elgon; one specimen.
_ A well-known species in South Africa and Madagascar.
ACHETID 2.
CURTILLIN A.
CURTILLA.
Gryllotalpa Latr. Hist. Nat. Crust. Ins. xi. p. 121 (1804), nom.
spec.
Curtilla Oken, Lehrb. Nat. iti. p. 445 (1815).
- 5. CuRTILLA AFRICANA.
_— Gryllotalpa africana Beauv. Ins. Afr. Amér. p. 229, pl. 2. f. 6
(1805).
Two specimens, 4000 feet, Baringo, Dec. 20, 1899.
A widely-distributed African and Kast-Indian species.
ACHETINE.
ACHETA.
Gryllus (Acheta) Linn. Syst. Nat. (ed. x.) i. p. 428 (1758).
Acheta Leach, Edinb. Encyel. ix. p. 119 (1815).
6. ACHETA BIMACULATA.
Gryllus bimaculatus De Geer, Mém. Ins. ii. p. 338, pl. 43. f. 1
(1773).
Acheta capensis Fabr. Syst. Ent. p. 281. n. 6 (1775).
Four specimens, Entebbe, Oct. 1900.
A common species throughout a great part cf Southern Europe
and Asia, and throughout all Africa.
PHASGONURID4.
MECOPODIN 2.
ANGDOPODA.
Anedopoda Karsch, Berl. ent. Zeitschr. xxxvi. pp. 333, 346 (1891).
7. ANGDOPODA LATIPENNIS.
Mecopoda latipennis Burm. Handb, Ent. i. p. 686. n. 2 (1838).
1902. ] BRITISH EAST AFRICA AND UGANDA. 95
Two specimens taken between Lake Victoria and Lake Tangan-
yika.
A common species in both Kast and West Africa.
HETRODINE.
ENYALIOPSIS.
Enyaliopsis Karsch, Berl. ent. Zeitschr. xxxi. p. 60 (1887).
8. ENYALIOPSIS PETERSII.
Hetrodes peters: Schaum, Ber. Akad. Ber]. 1853, p.777; Peters’s
Reise Mossamb., Zool. v. p. 119, pl. vii. f. 7 (1862).
Three specimens, from Mounts Elgon and Ruwenzori.
A common species throughout Kast Africa.
LOcUSTIDE.
TRYXALINA.
ACRIDA.
Gryllus (Acrida) Linn. Syst. Nat. (ed. x.) i. p. 427 (1758).
9. ACRIDA ACUMINATA.
Acrida acuminata Stal, Rec. Orth. p. 97 (1873).
Baringo, 4000 feet, Dec. 20, 1899.
Described from ‘“ Caffraria.” Three old specimens in the
Natural History Museum are from the ‘‘ Cape of Good Hope.”
PHLGOBA.
Phleoba Stal, Kugenie’s Resa, p. 340 (1862).
10. PHLG@OBA RUFESCENS, sp. Nn.
Male. Long. corp. 14 mm.; long. tegm. 9 mm.
Female. Long. corp. 21 mm.; long. tegm. 17 mm.
Rufo-testaceous ; antenne about 20-jointed, brown, except
towards the base, subensiform, broadest for about six of the
first joints of the flagellum and gradually tapering to the extre-
mity; head above with two obsolete reddish lines, marked with
a few black specks, diverging behind and continued still more
indistinctly on the pronotum. Pronotum at least twice as long
as broad; carinz pale yellow, the lateral ones edged below with
a black or reddish line: head with one or two indistinct reddish
(or, in the male, blackish) lateral stripes, continued on the sides of
the pronotum; these siles distinctly scabrous in the female, and
marked towards the front below the lateral carina with two large
smooth pits. On the upper surface the hinder lobe of the pro-
notum is punctured, the rest being nearly smooth; the hind
sulcus is placed behind the middle, and the two front ones are
widely interrupted in the middle, and only the second continued
96 MR. W. F. KIRBY ON ORTHOPTERA FROM [Feb. 4,
on the sides. Tegmina hyaline with reddish nervures; hind
tibie rather hairy, with eleven teeth on the outer edge.
Four specimens (1 ¢, 2 2, 1 nymph) from Baringo, Dee. 20,
1899.
There are several closely allied species from East Africa, of
which one only (P. alternata Schulthess, nec Brunner) has
hitherto been described.
PNOoRISA.
Gomphocerus (Pnorisa) Stal, Eugenie’s Resa, p. 341 (1860).
11. PnorisA CAPENSIS.
Stenobothrus capensis Walk. Cat. Derm. Salt. B.M. ii, p. 764.
n. 62 (1870).
Fifteen specimens, Baringo, Dec. 20, 1901.
None of these specimens exactly agree with the unique type of
the species, which is, besides, in rather poor condition ;. but they
vary so much among themselves that I cannot consider them
distinct. The males have indefinite longitudinal dusky markings
on the head and thorax (chiefly on the sides) as in Walker's type ;
but the latter has the outer central area of the femora much
more completely filled. up with blackish than in any of the
Baringo males, which have only isolated and variable blackish
patches on that area, the largest towards the extremity. In the
female there is a broad blackish band running backwards from
the eye, over the sides of the pronotum, and a portion of the
tegmina, and more or less distinctly bordered above and below by
yellow lines. The central area of the hind femora is filled up
externally in its upper half with a black stripe, broken into three
parts, and is bordered on the upperside above by two longer and
narrower black stripes.
LocusTIN&.
CHLGBORA.
Chlebora Sauss. Mém. Soc. Genéve, xxvill. (9) pp. 54, 132
(1884) ; xxx. (1) pp. 18, 19, 33 (1888).
12. CHLGBORA THALASSINA, Sp. 0.
Exp. al. 80 mm.; long. corp. 36 mm.
Female. Rufo-testaceous ; carina of the pronotum very slightly
raised, subobsolete behind, hinder half set with large granules,
and shorter and more rounded behind than in Humbe tenwicornis
Schaum. Tegmina with two oblique brown bands, the first at
one-quarter of the length, and the second, much narrower and
less complete, about the middle, bounding the outer subhyaline
area. Costal area spotted with brown, especially about the
first brown band. Beyond the lower extremity of the narrow
outer band a series of linear blackish marks runs along the
inner margin. Wings greenish yellow (possibly bright yellow in
1902. ] BRITISH EAST AFRICA AND UGANDA. 97
perfectly fresh specimens) towards the base, followed by a broad,
curved black band as in. Gastrimargus mar moratus, extending to
the anal angle and nearly touching the hind margin beyond-+the
curve ; apex of the wing hyaline.
One specimen, Mount Ruwenzori.
Closely allied to C. kellert Schulth., from Somali, but the pro-
notum is broader, less strongly ar ched and car inated, and more
rounded behind, and the black band on the hind wings is broader
and more regular.
GASTRIMARGUS.
Gastrimargus Sauss. Mém. Soc. Genéve, xxvill. (9) pp. 109,
110 (1884) ; xxx. (1) p. 37 (1888).
13. GASTRIMARGUS MARMORATUS.
Gryllus marmoratus, var. 3, Thunb. Mém. Acad. Pétersh. v.
p. 232 (1815); ix. p. 410, pl. 14. f. 3 (1824).
Pachytylus (Hdaleus) marmoratus Stal, Rec. Orth. i. p. 123
(1873)
Gdaleus marmoratus Sauss. Mém. Soc. Geneve, xxvii. (9)
p. 112, n. 2 (1884); xxx. (1) p. 39, n. 3 (1888).
One specimen of var. africana Sauss. (Mém. Soc. Geneve, xxx.
(1) p. 39) between Lakes Victoria and Tanganyika.
14. GASTRIMARGUS DETERMINATUS.
Pachytylus determinatus Walk, Cat. Derm. Salt. B.M. v.
Suppl. p. 72 (1871).
daleus verticalis Sauss. Mém. Soc. Genéve, xxviii. (9) p. 111
(1884).
Twelve specimens, Baringo, 4000 feet.
There are specimens in the Natural History Museum from the
Cape, Knysna, Natal, and Marabastaat.
PHYMATINA.
PHYMATEUS.
Phymateus Thunb. Mém, Acad. Pétersb. v. p. 257 (1815).
15, PHYMATEUS HGROTUS.
Pecilocera eyrota Gerst. Arch. f. Nat. xxxv. p. 216 (1869).
Three discoloured specimens, taken between Lake Victoria and
Lake Tanganyika.
Ac common species throughout Hast Africa,
DrcrvopHoRi#.
Feb testes TAPHRONOTA.
Taphronota Stal, Gifv. Vet.-Akad. Forh. xxix. p, 51 (1873).
16. TAPHRONOTA GABUNICA.
Taphronota gabunica K Karsoh, Ent. Nachr. XV. -p. 358 (1888).
Proc. Zoou. Soc.—1902, Vou... No. VII. 7
98 MR. W. F. KIRBY ON ORTHOPTERA FROM [ Feb. 4,
M nt Ruweuzori.
Two discoloured specimens, apparently belonging to this West-
African species.
DICTYOPHORUS.
Dictyophorus Thunb. Mém. Acad. Pétersb. v. p. 258 (1815).
|| Petasia Serv. Ann. Sci. Nat. xxii. p. 278 (1831).
17. DicTYOPHORUS ANCHIETA.
Petasia anchiete Bolivar, Jorn. Sci. Lisb. xxx. p. 110 (1882).
Three specimens from Entebbe, Oct. 1900.
Originally described from Angola, but apparently common
in East Africa. There are specimens in the Natural History
Museum from Abyssinia, Zomba, British East Africa, and
Tanganyika.
PAMPHAGINA.
XIPHICERA.
Xiphicera Lamarck, Anim. sans Vert. iv. p. 243 (1817).
18, XIPHICERA GIBBA, sp. 0.
Long. corp. 37 millim.; long. pron. 17 millim.; long. fem.
post. 124 millim. ; lat. 5 millim.
Male. Dark red ; vertex longer than broad, granulated, sloping,
and projecting in a point scarcely beyond the level of the lower
part of the face, to which it slopes gradually down; antenne,
funiculus apparently 6-jointed (the three basal joints hardly
separable), followed by a narrower joint but broader than long,
two intermediate joints, the first shorter than the other, and the
flagellum, which consists of only two joints, one longer than
broad, narrowed at. the base and truncated at the end, and the
terminal joint linear, Pronotum very high, laterally com-
pressed, granulated, pointed in front and bifid behind; in front
it slopes upwards to a rounded-off obtuse angle before the middle,
and then runs backwards to the extremity almost straight, the
hinder part being obtusely denticulated. Tegmina and wings
rudimentary ; back of abdomen with small teeth on the median
line, Hind femora granulated with white, and with the upper
surface straight and serrated, the extremity truncated ; the lower
edge moderately broadly laminated, denticulated, and with a
concavity before the extremity; hind tibie with strong short
spines, eight on the outer edge.
A single specimen from between Lake Victoria and Lake
Tanganyika, It would probably have fully-developed wings, if
mature.
Allied to XY, spinulosa Saussure and X, haploscelis Schaum.
MESAMBRIINA.
Two immature specimens from Baringo (Dec. 20, 1899),
probably belonging to a new genus allied to Mlesambria Stal.
Srey
1902.] BRITISH EAST AFRICA AND UGANDA. 99
CYRTACANTHACRINA.
CYRTACANTHACRIS.
Cyr Vacnittdiont is Walk. Cat. Derm. Salt. B.M. iii. p. 550 (1870).
19. CyRTACANTHACRIS PALLIDICORNIS, 0. 0.
Acridium ruticorne Burm. (nec Fabr.) Handb, Ent. ii. p. 630.
n, 9 (1838); Stal, Rec. Orth. i. p. 60. n. 2 (1873).
Acridiwm succinctum Serv. (nec Linn.) Ins. Orth. p. 642 (1839).
Seven specimens, between Lake Victoria and Lake Tanganyika.
There is also an immature specimen in the collection, perhaps
belonging to the same ; species, from Ruwenzori. This insect much
resembles Aeridiwm tataricwm Stal (nec Linn., which probably =
Schistocerca peregrina Oliv.). .
CATANTOPINA.
CATANTOPS,
Catantops Schaum. Monatsb. Berl. Akad. 1853, p. 779.
20. CATANTOPS CAPICOLA.
Acrydium (Catantops) capicola Stil, Kugenie’s Resa, p. 331
(1860).
Catantops humeralis Stal (nec Thunb.), Rec. Orth. i. p. 69. n. 1
(1873).
Baringo, 4000 feet, Dec. 20, 1901.
A rather large specimen, measuring 48 millim, in expanse.
CALLIPTAMINE.
EuryPHYMUS.
ELuryphymus Stal, Rec, Orth. 1. p. 72 (1873).
21. HURYPHYMUS CRASSUS.
Oaloptenus crassus Walk. Cat. Derm. Salt. iv. p. 694. n. 39
(1870).
O. llepidus Walk. |.c. n. 40 (1870).
Var. C. pinguis Walk. 1. c. n, 41 (1870).
Twenty specimens from Baringo, 4000 feet, Dec. 20, 1899.
Walker’s specimens are from Natal, so far as they were labelled
with any special locality.
EUPREPOCNEMIN#.
HETERAGRIS.
Heteracris Walk. Cat. Derm, Salt. B. M. iv. p. 655 (1870).-
Demodocus Stal, Bihang Vet.-Akad. Handl. v. (4) p. 75 (1878).
As S$tal’s name Demodocus is preoccupied in Coleoptera, I
propose to restrict Walker’s name Heteracris to this genus,
100 MR. W. F. KIRBY ON ORTHOPTERA FROM [ Feb. 4,
22. HETERACRIS BETTONI, sp. n.
Male. Long. corp. 31-40 millim. ; long. al. ant. 14-15 millim. ;
long: fem. post. 16-17-millim.
Female. Long. corp. 31-40 millim.; long. al. ant. 20-34 millim. ;
long. fem. post. 25-33 millim.
~ Chestnut-red, lighter in the male than in the female; vertex
much contracted between the eyes, with a shallow depression
triangularly expanded on each side beyond the contraction: this
is intersected by a slight carina, the continuation of the middle
carina of the pronotum. Pronotum with the hinder lobe granu-
lated ; in front of this portion extends a brown shade, divided by
the central carina, and narrowing in front, but filling up most of
the centre of the pronotum and vertex as far as the end of the
depression on the latter. Head finely punctured ; sides of the
pronotum more sparingly but more coarsely punctured than the
hinder part, and with two very large pits below the lateral
carina in front of the first suture, and behind these two others
between the two sutures in the female, but only a still larger
one in the male; below this, on the middle of the central lobe of
the pronotum, is an oblong yellow carina. Legs red, first and
second pairs short, hind legs very long, femora with the basal half
moderately thickened ; tibize about as long as the femora, with
from twelve to fourteen spines, slightly black-tipped, and gradu-
ally increasing in length from the base to the extremity. First
and third joints of tarsi of about equal length, the first thickened
in the middle, especially beneath ; the second about three-fifths as
long. Tegmina brownish hyaline, with dark nervures and the
outlines of several large spots ; some of the longitudinal nervures
and intermediate spaces are reddish, especially above and below
the central area; in closed specimens the lower red stripe is seen
to be continuous with the red borders of the pronotum. Wings
clear hyaline, with brown and reddish nervures.
Described from three males and one female from Baringo
(Johnston); one female from Maungu, B. EK. Africa (Betton) ; one
female from Thika-Thika, B. E. Africa (Gregory); one female
from Mombasa (Dr. J. Wilson); one female from Atbara,
Abyssinia (purchased).
The female from Baringo is smaller than any of the others.
Apparently allied to Heteracris speciosa Walk. from Sierra
‘Leone, of which the Museum at present possesses only the type,
an immature specimen.
CALOPTENOPSIS.
Caloptenopsis Bolivar, Jorn. Sci. Lisb. (2) i. p. 173 (1889).
Head with the vertex between the eyes narrowed and tri-
carinate, the carine very short, and formed by an oblong fovea on
each side of the central carina; beyond this point the vertex
slopes smoothly into the broad frontal ridge, which is not
‘sulcated but sparingly punctured, and is nearly straight. Thorax
broad, tricarinate, depressed, with the central carina higher than
1902.) BRITISH EAST AFRICA AND UGANDA, 101
the others; principal suture straight, a little in front of the
middle; second suture curved backwards above and diverging
from the hinder one on the sides ; front suture scarcely continued
on the sides, but there is a lateral suture near the front of the
pleura, curving backwards below. Pectoral tubercle large and
thick, obtusely rounded, broader than long. Pleura long, pro-
jecting a little on the middle edge below, and rounded. Hind
temora thick, not much narrowed towards the extremity and
denticulated above. Hind tibie with- six outer and eight inner
spines, and provided with five terminal spines, the middle one
inferior, as long as the metatarsus, hairy, with a strong tooth on
the upper surface before the extremity. Metatarsus trilobate
beneath ; second joint short above, but with a projection below
the base of the terminal joint; the latter is slender, with large
claws and pulvillus, and a little shorter than the metatarsus.
Male with the cerci very broad, somewhat spatulate at the
extremity, and furnished with a short black terminal tooth on
the outer side. Female with the upper appendages longer than
the lower ones and hooked upwards at the extremity ; the lower
appendages hooked downwards.
A remarkable and interesting genus, which I had characterized
as new before recognizing its identity with Caloptenopsis ; and
as my definition was already in print, I have allowed it to stand.
My definition, of course, is taken from C. johnston.
There are now several species of Caloptenopsis known, chiefly
from Kast and Central Africa.
23, CALOPTENOPSIS JOHNSTONI, sp. n.
Long. corp. ¢ 17, 2 21 millim.; exp. al. ¢ 25, 2 40 millim,
Rufo-testaceous ; face not carinated, clypeus broad below, with
a black spot.at each lower angle. Pronotum varied with blackish
in the middle above, especially in front; the inner borders of the
lateral carine closely punctured; the hinder lobe and the meta-
pleura thickly and closely punctured. Front of the pleura, below
the lateral carine, with two large pits, the first double, below
these is a brown curved band, ceasing at the hinder suture;
below it is a second but shorter brown stripe. The colour of the
pleura is paler than the upper part of the pronotum. Subalary
spaces with large punctures. Both surfaces of the hind femora
with curved or slightly angulated black lines, followed by black
dots on the carine; the spaces between the outer and middle
carins above and between the three lower carine unspotted.
Upper curve of the knees black on both sides, space below this
yellowish. Tibial spines rather small and tipped with black.
Wings brownish hyaline, interspersed with brown network-
patterns in the middle, becoming more macular towards the
extremity ; anal area with reddish nervures.
Described from three males and four females from Baringo,
Dee, 20, 1899.
102 DR. C, I. FORSYTH MAJOR ON PLIOCENE VOLES, [ Feb. 18,
February 18, 1902.
Prof. G. B. Howss, LL.D., F.R.S., Vice-President,
in the Chair.
Mr. L. W. Byrne, F.Z.8., called the attention of the Meeting
to the description of Lepidogaster stictopteryx, a supposed new
species of Sucker-fish, which had been given by Mr. E. W. L. Holt
and himself in a communication made to the Society on
November 15th, 1898, and made the following remarks :—
On November 15th, 1898, we exhibited before this Society
(P. Z. S. 1898, p. 589) specimens of a Lepidogaster. We supposed
them to be attributable to a new species, for which we suggested
the name of L. stictopteryw. =~ Af ae
The examination of further specimens has convinced us that
we have been guilty of adding further confusion to the synonymy
of the species of this genus, and that our L. stictopteryx is not
specifically distinct from LZ. microcephalus Brook, the synonymy
of which should stand as follows :—
LEPIDOGASTER MICROCEPHALUS.
L. microcephalus Brook, Proc. Roy. Phys. Soc. Edin. x. p. 166,
pl. vii. (1888).
2L. bimaculatus $, Guitel, Comptes Rendus, exl. p. 759 (1890).
L. stictopteryx Holt & Byrne, P. Z.8. 1898, p. 589.
Fortunately our friend Professor Guitel, of Rennes, is con-
tinuing his studies upon this genus, and informs us that he has
obtained at Roscoff material which he believes will enable him to
deal with the question in a satisfactory manner. Under these
circumstances we feel it would be superfluous for us to do more
than correct our own mistake, and we have entrusted our notes,
drawings, and material to his most able hands.
Mr. W. B. Tegetmeier, F.Z.S., exhibited and made remarks upon
the skull of a supposed hybrid between the Sheep and the Pig,
named “Cuino”?* by the inhabitants of Mexico, where it was
extensively reared as anagriculturalanimal. The skull was clearly
that of a Pig.
Dr. C. I. Forsyth Major, F.Z.8., exhibited some Jaws and
teeth of Pliocene Voles (Zimomys, gen. nov.), from the Norwich
Crag at Thorpe, and from the Upper Val d’Arno; and made the
following remarks :—
The Pliocene remains of Voles here exhibited—a mandibular
ramus from the lacustrine beds of the Upper Val d’Arno in
Italy, and over forty bits from the Norwich Crag at Thorpe,
1 For information respecting this supposed hybrid see ‘ Field,’ vol. xevi. (1900)
p. 497, and xevii. (1901) p. 233.
1902. | DR. C. I. FORSYTH MAJOR ON PLIOCENE VOLES. 103
mostly isolated teeth—are so minute and fragmentary, that I have
to supplement my demonstration by sketches.
The jaw from the Val d’Arno (text-fig. 13, nos. 8, 9), con-
taining two anterior rooted molars, was mentioned by me
upwards of twenty years ago. The first lower molar (text-
fig. 13, no, 8) exhibits in its anterior portion an enamel islet,
which is a very strange feature ina Vole’s molar. I am sorry
to trouble members with such a minute detail; but almost the
whole interest centres around this insular eccentricity, so to
Text-fig. 13.
H. G. del.
Teeth and jaws of Tertiary Voles.
Figs. 1-5 & 7 represent the first lower molars, upper view.—Fig. 1. Mimomys
intermedius (Newt.), West Runton Forest Bed (B. M. No. 6968 d, from Savin
Coll. No. 1705): left side—Figs. 2 & 3. Mimomys pliocenicus (Maj.),
Norwich Crag, Thorpe (Norw. Castle Mus. No. 971): fig. 2, left side; fig, 3,
right side.—Figs. 4&5. Mimomys pliocenicus (Maj.), Norwich Crag, Thorpe
(Norw. Castle Mus. No. 551, from Fitch Coll.) : fig. 4, right side; fig. 5, left
side (figured by E. T. Newton, ‘ Forest Bed,’ pl. 13. fig. 13).—Fig. 6. Mimomys
pliocenicus. (Maj.), third upper molar, left side; Hast Runton Forest Bed
(B. M. No. 6967, from Savin Coll. No. 464),.—Fig. 7. Mimomys newtoni, sp. n.,
Hast Runton Forest Bed (B. M. No. 6967 a, from Savin Coll. No. 480) : left
side —Fig. 8. Mimomys pliocenicus (Maj.), first and second lower molars,
upper view ; Upper Val d’Arno, Italy (Florence Museum).—Fig. 9 a. The same
specimen, outer view of the mandible, nat. size.—Fig. 9 6. The same enlarged,
r=root of m?,—Fig. 9 c. The same mandible, inner view, enlarged.
104 DR. C, I. FORSYTH MAJOR ON PLIOCENE VOLES. [Feb, 18,
speak, of the fossil tooth. I had before met with a similar
feature in one of two very young teeth of the recent amphibius-
group, from Pisa, which presumably belonged to Savi’s Arvicola
destructor. Inthe recent tooth the enamel islet showed a slightly
different position and genesis, was quite superficial and therefore
ephemeral; it was associated with some other complications—
two additional shallow enamel-loops—which likewise approach
the tooth of the very young amphibius to that of the Pliocene
form. In the recent species this pattern is very soon worn away ;
there is no more trace of it in slightly older specimens. This is a
fresh instance of a recent form preserving in the younger stages
of its molar the features of a Tertiary form.
When during the revision of the fossil Rodentia of the British
Museum, the Microtide of the Forest Bed came to be studied, I
was anxious to ascertain whether the enamel islet occurred there
too in adult specimens, as is the case in the Val d’Arno fossil.
Mr. E. T. Newton has published an elaborate description of
the Rodents of the Forest Bed and Norwich Crag, and has shown
that the larger Voles are, by the presence of well-developed fangs to
their molars, very distinct from the amphibius-type with which they
had been confused by all previous writers. He who enjoys the
advantage of standing on his predecessor's solid shoulders, has also
the duty to try and see a little farther, especially when additional
material has accumulated in the meantime. If, therefore, to-day
a step forward is possible in the knowledge of the Pliocene Voles,
it is but fair to acknowledge that this is in a great measure due
to Mr. Newton’s previous work.
Among the Voles’ teeth of the Savin Collection from the
Forest Bed, I found the character alluded to, but only in a
relatively small number of teeth (e.g. text-fig. 13, no. 1) and in
different proportions according to the localities. Whilst among
55 first molars from the West Runton Upper Freshwater Beds
only four showed the character in question, the number of teeth
provided with enamel islets was larger among the less numerous
Voles’ remains from the East Runton Forest Bed, and, more-
over, other features became apparent.
From the Norwich Castle Museum I have received of late,
through the kindness of Mr. Leney, a small number of teeth and
two jaws, here exhibited, which were collected by Mr. Fitch in
the Norwich Crag at Thorpe (text-figs. 13, nos. 2-5; 14, no. 15;
15, nos. 20, 29). Here the presence of the enamel islet is the
rule: there are ten anterior lower molars in this small series—
eight exhibit the islet, one is very old and apparently has lost
evcry trace of it; the tenth, a very young tooth (text-fig. 15,
no. 29), reveals the genesis of the islet, which is the central
portion of the antero-external enamel fold. Moreover, the teeth
are of two different sizes.
In short, the result of the investigation is, that the Voles of
the Norwich Crag are different from those of West Runton and
are represented by two species; whereas at East Runton the
1902. ] DR, C, I, FORSYTH MAJOR ON PLIOCENE VOLES. 105°
West Runton type occurs together with the Crag types. The
larger of the Crag-forms is besides represented by specimens from
Bramerton (text-fig. 14, nos. 14a, 6) and from Kyson in Suffolk.
Text-fig. 14.
Teeth of Tertiary Voles, enlarged.
Figs. 10-17. Views of first lower molars. a==outer aspect, 6=inner aspect (except
146=upper view).—Fig. 10. Wimomys newtoni, sp. n., same specimen as
fig. 7.—Fig. 11. Mimomys intermedius (Newt.), same specimen as fig. 1.—
Fig. 12. Mimomgs intermedius (Newt.), same specimen as fig. 21.—Fig. 13.
“ Mimomys intermedius (Newt.),” same specimen as fig. 26, right side; West
Runton (B. M. No. 6968 e, from Savin Coll. No. 1705).—Fig. 14a. Mimomys
pliocenicus (Maj.), Norwich Crag, Bramerton (Norwich Castle Mus. No. 728,
from Reeve Coll.).—Fig. 146. The same, upper view.—Fig. 15. Same speci-
men as fig. 20, Mimomys pliocenicus (Maj.), Norwich Crag, Thorpe (Norwich
Castle Mus. No. 971).—Fig. 16. Mimomys newtoni (?), Hast Runton Forest
Bed (B. M. No. 6967 6, from Savin Coll. No. 464).—Fig. 17. Wimomys plio-
cenicus (Maj.), same specimen as fig. 5.
For the present I content myself with three specific names,
calling (1) pliocenicus the larger Crag-form, which I identify
with the one from the Val d’Arno; (2) newtoni, a smaller rooted
106
DR. C, I. FORSYTH MAJOR ON PLIOCENE VOLES. [{ Feb. 18,
Text-fig. 15.
Teeth of Voles from Forest Bed and Norwich Crag.
Figs. 18-21, 22 a, 23-80, first lower molars, upper view, enlarged.
Fig. 18. Microtus, sp., recalling somewhat IM. gregalis ; rootless, left side; West
Runton (B. M. No. 6987 a, from Savin Coll. No. 1708).—Fig. 19. Microtus
nivaloides, sp. n., recalling WM. nivalis, but smaller and anterior loop more
produced : left side; rootless (B. M. No. 6987 6, from Savin Coll. No. 1708).
—Fig. 20. Minomys pliocenicus, same specimen as fig. 15: left side, enamel
islet vanishing.—Fig. 21. Mimomys intermedius, same specimen as fig. 12 ;
West Runton (B. M. No. M 6968 a, from Savin Coll. No. 1692): left side,
cement-spaces closed below, but roots not yet developed.—Fig. 22. Mimomys
intermedius, first lower molar, right side, outer aspect; West Runton (B. M.
No. 6968 8, from Savin Coll. No. 1692) : cement-spaces closed below, but roots
not yet developed; fig. 22a. Same specimen, upper view; fig. 226. Same
specimen, inner aspect.—Fig. 23. Mimomys newtoni?, Hast Runton; same
specimen as fig. 16.—Fig. 24. Mimomys intermedius, right side; Kast Runton
(B. M. No. 6967 ¢, from Savin Coll. No. 465): cement-spaces, with the ex-
ception of the antero-external, still open below.—Fig. 25. Mimomys inter-
medius, right side, young ; West Runton (B. M. No. M6968 ec, from Savin Coll.
No. 1692): cement-spaces narrowing below, but not closed (the lower end of
the tooth is incomplete from break). From an inspection of the outer side
it becomes evident that by progress of wear the anterior enamel fold would
soon have been reduced to an ephemeral enamel islet.—Fig. 26. Mimomys
intermedius ?, same specimen as fig. 13, right side; West Runton.—Fig. 27.
Wicrotus sp., recalling WM. arvalis: left side, rootless ; West Runton (B. M.
No. M 6987 ¢, from Savin Coll. No, 1708).—Fig. 28. Microtus (Pitymys) sp.,
right side, rootless; West Runton (B. M. No. M 6987 d, from Savin Coll.
No. 1708). Behind the anterior loop follows a transverse loop resulting from
the union of the third outer and the fourth inner prism (counting from
behind). This loop is separated from the anterior loop by the meeting in the
middle line of the third outer and the fourth inner reentrant angle; features
characteristic of the European members of the subgenus Pitymys.—Fig. 29.
Mimomys pliocenicus, young, right side, posterior portion broken off; Norwich
Crag, Thorpe (Norw. Castle Mus. No. 971: Fitch Collection). The antero-
external reentrant angle not yet reduced to an enamel islet. Cement-spaces
beginning to be closed below.—Fig. 30. Mimomys pliocenicus, young, right
side (Norwich Mus. No. 2708: from Gurney Collection). ‘“ Upper Fresh-
water Bed, Ostend.” Cement-spaces closed below; no roots developed.
Enamel islet not yet formed.
1902. ] MR. R. LYDEKKER ON AN ELK FROM SIBERIA, 107
form from the Norwich Crag and East Runton (text-fig. 13,
no. 7), which has characters of its own; (3) Mr. Newton’s name
intermedius is restricted to the form or forms met with at West
Runton (text-figs. 13, no. 1; 14, nos. 11-13; 15, nos. 21, 22, 25,
26) and a few from East Runton (text-fig. 15, no. 24). Iam,
however, quite convinced that at least double this number of
species ought to be recognized, and am only prevented from doing
so at present because I do not wish to found species on isolated
teeth.
The larger Crag species is mainly characterized by the presence
of the enamel islet, except in quite old specimens ; by the earlier
development of fangs; by the presence of an enamel islet in the
last upper molar also; and by the presence of three roots in the two
anterior upper molars, whereas in J. intermedius these same
teeth have only two roots.
The four anterior lower molars of MW. intermedius, in which the
islet occurs, are all very little worn, although full-grown. It is
therefore probable that the enamel islet will be found to be a
constant: feature in the young teeth of J. intermedius, but more
ephemeral than in J. pliocenicus. I propose to form a distinct
genus, Mimomys, for all these Voles with rooted molars, which
are clearly different from Hotomys, Phenacomys, and Dolomys.
M. newtoni may prove, hereafter, to form a distinct genus.
In the Savin Collection there are, besides those already referred
to, about 17 fragmentary jaws and a small number of isolated
teeth of JMicrotus, provided with rootless teeth throughout life ;
with one exception, a single anterior lower molar tooth from East
Runton, they all came from the West Runton Upper Freshwater
Beds. There are at least four different forms; several of them
show in the conformation of their lower anterior molar some
resemblance to such living forms as MW. arvalis (text-fig. 15, no. 27),
M. nivalis (text-fig. 15, no. 19), and I, gregalis (text-fig. 15, no. 18),
On closer inspection I find, however, that, with the single exception
of one isolated tooth, I can refer none of these remains to any
recent nor to any hitherto known fossil species. The majority of
the rami, ten in number, have nothing to do with the subgenus
Microtus, but show in their anterior lower molar a feature (text-
fig. 15, no. 28) which is characteristic of the Kuropean and some
of the American members of Pitymys, and is found almost
identical in the North-American Pedomys.
Mr. Lydekker exhibited the skull and antlers (text-fig. 16,
p. 108) of an adult male Elk from Siberia, together with the
antlers of a second example, lent by Mr. Rowland Ward.
The skull indicated an animal of at least 6 or 7 years old,
the cranial sutures being for the most part obliterated, while the
cheek-teeth were about half-worn. It was that of a somewhat older
animal than the one to which an American skeleton mounted in
the British Museum belonged. In the latter the palmation of the
108 MR. R. LYDEKKER ON AN ELK FROM SIBERIA. __[ Feb. 18,
antlers was well developed ; but in both the pairs of antlers ex-
hibited there was practically no palmation. These antlers were in
fact very like those of young Scandinavian Elk, only with the
palmation still less. They showed three tines on each side on the
upper or hinder half of the main bifurcation, and either one or
two tines on the lower or front branch. Mr. Lydekker had been
informed that other Elk-antlers from Siberia were of a similar
type. :
Text-fig. 16.
Skull and antlers, with the upper cheek-dentition (a), of Siberian Elk.
From the type specimen in the Museum at Tring Park,
That the specimens exhibited were not the result of senile
decadence was quite evident, not only from the symmetrical form
of the antlers themselves, but likewise from the state of wear of
the cheek-teeth (text-fig. 16, a) of the skull,
The similarity of the two pairs of antlers, together with the
information as to this type being characteristic of all Elk-antlers
from the same country, induced Mr. Lydekker to regard the Elk
of Siberia as a distinct form. Whether it should be considered a
species or a variety was a somewhat difficult question ; but since
the antlers exhibited involved a modification in the definition of
the genus, it seemed advisable to allow specific rank in this case,
JE LoS ISN wold tel Wilk,
Le he
aa,
2) EDO
5
H.Grénvold,del. Photogravure by Bale &Danielsson.L'?
STU Iie (Ole MOOS Wis Ie yAy VAIL gave PIE LGA.
1902. } "ON MUSTELA PALHATTICA. 109
Although in a recently published work Mr. Lydekker had
suggested the possibility of the Siberian Elk proving distinct, so
far as he was aware it had not yet receiveda name. An Elk
with antlers not unlike those of the specimens exhibited had
been described in 1847 by Rouillier, in Fischer de Waldheim’s
‘Jubileum,’ under the name of Alces resupinatus, based on a
skull from a Pleistocene deposit in Russia. There did not appear,
however, to be any characters by which that specimen could be
distinguished from young skulls of the Scandinavian Elk.
Under these circumstances Mr. Lydekker proposed to name
the Siberian Elk Alces bedfordie, in honour of the wife of the
President of the Society. This species would be distinguished
from both the Scandinavian and American races of Alces machlis
by its non-palmated antlers, which carried only four or five tines
on each side. The complete specimen exhibited would form the
type.
vthe occurrence in Siberia of an Elk with antlers of the simple
type of those exhibited was a fact of considerable interest, since
that country was probably the centre whence both the European
and American races of the true Elk were evolved.
[| P.S..—Since this exhibition took place Mr. Lydekker had seen
five other pairs of Elk-antlers from Siberia, all of the same form.
Three of these specimens, together with the two exhibited, had
been acquired by Mr. Walter Rothschild.
The following papers were read :—
1. On Mustela paleattica from the Upper Miocene of
Pikermi and Samos. By C. I. Forsyra Magor.
[Received December 17, 1901.]
(Plate VIL.)
The type of Weithofer’s Wustela paleattica’, from Pikermi, is
in the Vienna Museum. It is represented by a badly crushed
skull (of which, however, the teeth, minus the incisors, are very
well preserved), by the two almost intact mandibular rami, and
by part of the skeleton. The whole was kept together and
preserved from total destruction by being lodged between the
rami of a Hipparion mandible.
The characteristic features of this species are furnished by the
conformation of the upper and the talon of the anterior lower
molar. Whilst the posterior upper premolar (p.1) bears the
characteristic features of Mustela, in its elongate outlines and the
1 For explanation of the Plate, see p. 114.
* A. Weithofer, “Beitrage zur Kenntniss der Fauna von Pikermi bei Athen.”
(Beitr. Pal. Oesterreich-Ungarns, vi. pp. 226-231, pl.-x. figs. 1-11 (1888). ]
110 DR. C. I. FORSYTH MAJOR ON [ Feb. 18,
anteriorly situated small inner cusp, separated by a constriction
from the blade, the molar approaches somewhat JMeles by being
more extended antero-posteriorly than in the recent species of
Mustela proper, and by the presence of a third cusp to the inner
side of the postero-external tubercle. Likewise, the lower m. 1
has its talon more complicated than in Mustela.
In the same year as Weithofer’s publication I identified with
his species an incomplete skull (the facial part missing) ’ which I
had discovered in the contemporaneous deposits of Samos, in the
locality Andriano near the village Mitylini.
Schlosser” discusses Weithofer’s type under the heading
“ Meles? (Mustela) paleatiica,’ and unites with it an isolated
upper molar, also from Pikermi, which he had formerly been
disposed to regard as “‘ Martes pentelict Gaudry,” and which there-
fore bears this name in the explanation of his plate viii. (fig. 16).
Two years later® Weithofer’s type was registered by Schlosser as
Meles paleatticus.
Von Zittel in his turn proposes for it the new generic name
Promeles, and places it with the Melinw*; whereas Winge refers
Promeles to the Mustelinw, and places it side by side with
“ Martes”°.
In the Geological Museum of Turin I came upon the skull of a
small Carnivore from Pikermi, which had been received many
years before. It was kindly intrusted to me by Professors Parona
and Sacco, and proved, when cleaned, to belong to the same
species as the one described by Weithofer. Being so much more
complete than my specimen from Samos, I have preferred to
describe and figure the Turin specimen rather than the latter.
The skull is somewhat laterally compressed and otherwise
distorted. Both zygomata are incomplete, the left one less than
the right. The principal lesion is in the lateral region of the
right side, the posterior part of the frontal and the parietal being
lost. The mandible was in its natural position, and so firmly
adhering to the skull, that to detach it as a whole would have
been impossible without endangering the teeth. I determined
therefore to sacrifice part of the right mandibular ramus—which
was already damaged—rather than spoil the teeth, and succeeded
in developing satisfactorily the m.1, p.1, and p.2 of the right
upper, and the m.2, m.1, and p.1 of the right lower jaw.
Weithofer assigns to his specimen the size of Mustela martes,
only slightly more robust; the teeth as figured show no signs of
wear. The Turin specimen shows the teeth moderately worn, and,
as will appear from the measurements, it was slightly larger than
1 @. Rendus, 31 Dec. 1888, p. 1179. It is no. 272 of Mr. W. Barbey’s collection at
Valleyres (Switzerland). See Forsyth Major, ‘Le Gisement ossifére de Mitylini et
Catalogue d’Ossements fossiles recueillis 4 Mitylini, Ile de Samos,’ p. 27, no. 272
1894).
2 Pal. Oesterr.-Ung. viii. p. 352 (1888).
3 Td. op. cit. viii. p. 469 (1890).
4 Handbuch der Palaeont. iv. p. 690 (1890).
5 E Museo Lundi, (2) iv. pp. 66, 69 (1895).
1902. ] MUSTELA PALHATTICA, Lid
the former. On comparing Weithofer’s figure 1 (pl. x.) with my
figures (Pl. VIT.), the difference in size would appear still greater ;
but I find that the size of his figure does not in every respect
correspond with the measurements given in the text, whereas, on
the other hand, the statement on p. 228, that the upper series of
cheek-teeth have together a length of 34 millim., is obviously a
misprint: presumably we have to read 24 millim.
Turin. Vienna.
mm. mim.
Greatest length of upper molar (m.1) ... 75 6°5
“3 breadth ‘ x PEF) ROS 10-0
“3 length of upper carnassial (p. 1) 9-2 9-0
breadth of p.1 underneath
principal cusp eeea.2 eee se 4:0 37
Mensti of Upper Ya: ssc) ease aaestece nee 6:0 6-0?
35 Ah ois eee Ae AAR habs 4:3 4:0°
ue a BD APL, ES Se etnt eaOne 2°5
Length of the cheek-teeth series in the Turin specimen, from
posterior border of third cusp of m. 1 to anterior margin of p. 3,
25°5 millim.
Turin. Vienna.
Length of mandibular ramus (from mm. mm.
anterior basis of canine to condyloid
[ORCOSSS)) CouagesanccusosonsescussnoodgcesucS. 57:0 56°5
Height of coronoid process ............045 29:0 29-0
From lower margin of foramen magnum
to anterior basis of Incisors ............ ca. 79-0
From hinder margin of teeth-series to
posterior palatal emargination ......... 10°0
engthrofllowerner2 .1.)).is.voresscselsadees
e ee DAO REE SES AIS ED ese 12:4 11:8
3 9 UNG AT on ce Senne ae eee 6°5 6:3
53 , V3 ho ison oe Soda baat ign Oa 5:0 4:8
sy 2 Tip Gabe o Sona san at Soadoonn oad: 45 4:3
3 Ag canines at basis ......... 57
The six upper incisors are in place in the Turin specimen
(Pl. VII. fig. 3); the anterior portion of the mandible remaining
attached to the skull, only their anterior side is visible. They
correspond perfectly with the incisors of d/ustela, without any
trace of the tricuspid condition of the blades exhibited by the
upper incisors of Meles, Mydaus, Helictis, Mephitis, and Conepatus.
The upper canines are both broken, only the stumps remaining.
Weithofer describes the upper canines of the Vienna specimen as
being “etwas hakig nach riickwirts gebogen.” In the Meline
the upper canines are more in the shape of a dagger, the backward
1 The tooth of the Turin specimen being worn, the measurement of height is
omitted. : : :
2 Weithofer terms this tooth y. 1, 8 Weithofer, sub y. 2.
112 DR. C. I. FORSYTH MAJOR ON [Feb. 18,
‘curvature of the Musteline canines being absent or scarcely
appreciable. eo
The anterior premolar (p.4), absent in the Vienna. skull, is
present on both sides in the Turin specimen. I have nothing to
add to Weithofer’s description of the two following premolars
(p. 2 and p. 3).
The perfectly Musteline character of the upper carnassial (p. 1)
has already been mentioned. The only difference, already pointed
out by the Austrian paleontologist, from recent species of Mustela
is in the outer contour, which is almost convex in the fossil,
slightly concave in recent species. The inner cusp is less con-
stricted than in UZ. martes, M. zibellina, and M. pennanti, but
resembles in this respect the MW. foina.
Of the upper molar (m.1) Weithofer says that it is different
from that of all the other Mustelide, and he describes it minutely as
follows! :—‘Er ist bedeutend stirker entwickelt, mehr complicirt
in der Richtung gegen den Dachs hin, ist iiberhaupt nur ein
verkiirzter Dachszahn mit allen den Elementen, die diesen charak-
terisiren. Die beiden iiusseren Tuberkel des Marderzahnes sind
viel stiirker, stehen in ihrer Entwicklung in der Mitte zwischen
Marder und Dachs und iiberdies ist bereits auch der dritte dussere
Tuberkel des Dachszahnes vorhanden. Von diesem zieht sich
eine hockerige, in zahlreiche kleine Tuberkel aufgeléste Wulst
gegen innen, und, an der Innenseite das Zahnes, gegen vorne,
welche in dieser Weise ebenfalls nur beim Dachs auftritt, noch
nicht aber beim Marder. Zum Unterschiede von ersterem theilt
sie sich jedoch in ihrem Verlaufe an der Innenseite riickwirts in
zwei Aeste, welche beide die erwihnte grobe Kérnelung besitzen.
Der beim Dachs in der Mitte dieses Zahnes auftretende, von der
Vorderecke ausgehende Kamm, der sich meist in drei Hocker
auflést und dessen Aequivalent beim Marder nur ein einfacher
kleiner Tuberkel ist, ist hier auch als ziemlich langer, bogenformig
gekriimmter Kamm ausgebildet. Die Gesammtform des Zahnes
ist eine mehr parallelopipedische, wenigstens ist die Vorder- und
Hinterkante vollstindig gleichlaufend, welche beim Marderzahn
nach aussen stark convergiren. Es ergibt sich daraus eine
besondere Ausdehnung des Aussenrandes, waihrend ber Innenrand
nur wenig grésser ist als beim Marder.”
On comparing this minute description and the figure of the
tooth (pl. x. fig. 1) with the specimen at my disposal (Pl. VII.
figs. 3 & 4), we have to bear in mind that the teeth of the Vienna
specimen are scarcely touched by wear, and that the granulations
-of the talon, of which there are only traces remaining in the worn
-tooth of the Turin specimen, are quite as conspicuous in unworn
molars of recent Mustelas as they are in the Vienna. tooth.
The general form of the molar is dumbbell-shaped in J. martes,
M. zibellina, M. pennanti, and M. foina. In Weithofer’s
‘specimen the outer and inner margins run perfectly parallel
1 Op. cit. p. 228.
1902.] MUSTELA PALHATTICA, 113
to each other, but in the Turin specimen there is a slight emargi-
nation on the posterior margin, internally from the third cusp ;
and the same may be seen in a fine skull of J. paleattica from
Pikermi, which forms part of Dr. A. Smith Woodward’s recent
successful excavations at Pikermi. In the Valleyres, Turin, and
London specimens the interior margin of the tooth is slightly
more elongate than the exterior, so that the anterior and the
posterior margins converge slightly towards the outer sides.
Conversely, in the Indian Martens (the WU. flavigula- group)
“this molar differs in form from that in IZ. foina and MW. martes
by having the inner lobe no broader from back to front than the
outer.” ?
On the whole the fossil tooth differs from the molars of the
martes-group by slight characters only. Now, the only reason for
collocating the fossil within the dJelinw has been the shape of
this upper m.1, with which of course goes hand in hand that of
the talon of the lower m.1l. It is, however, to be considered
that a tooth situated at the posterior end of the series is always
liable to vary more or less, and within the JM/eline in particular
this tooth is by no means characteristic for the group. In
Helictis it is quite narrow, the inner part not broader than the
outer and the anterior and posterior margin almost parallel. In
Meles the outer margin is much shorter than the inner, so that in
this respect J/eles agrees more with Mustela martes and allied
species. In Mephitis and allies there is no trace of a third cusp,
which, on the other hand, is present, although feebly developed, in
the unworn molars of some species of Justela (e. g., MZ. pennanti,
M. zibellina), as also in unworn specimens of the South-African
Mellivora (M. ratel), as well as in the Vison.
Far more characteristic is the upper carnassial (Pl. VII.
figs. 3 & 4). In the Musteline and in “ Promeles” it shows
the elongate form and the small anterior talon-cusp. The region
of the cheek is more sharply separated than in recent Mustela
from the nasal region by a blunt arcuate ridge, and in relation
with this the depression in front of the orbits and above the
infraorbital foramen is deeper.
The anterior and posterior roots of the zygoma rise almost
vertically, so that the zygoma is shown to have a highly arcuate
form, just as in Mustela.
The palate is more prolonged behind the molar series than in
the recent species.
The under contour of the mandible is slightly more arcuate in
the fossil than in WZ, zibellina and MW. martes, and even more than
in Mustela foina as described.
In all the Meline the carnassial is short and often provided
with two or three talon-cusps. When there is only one (Mephitis),
this is placed opposite the middle of the blade, and on either side
connected by a cingulum with the antero- and the postero-
1 W. T. Blanford, ‘ The Fauna of British India ’—Mammalia, p. 159 (1888).
Proc. Zoo. Soc,— 1902, Vou, I, No, VIII. 8
114 MR. OLDFIELD THOMAS ON [Feb, 18,
external margin of the blade, so that a valley is formed between.
the inner and the outer part of the tooth.
By. the conformation of its upper carnassial, therefore, the fossil
is excluded from the MWeline—the upper and the lower one making
only a slight approach towards the form they have im some
members of this subfamily ; whilst the characters of the skull and
of the skeleton, so far as known, bring it likewise in closer con-
nection with the Musteline and with Mustela in particular. To
emphasize this, it seems preferable to leave it in the latter genus,
viz., to revive the name by which it was originally described—
Mustela paleattica Weith.
EXPLANATION OF PLATE VII.
Skull of Mustela paleattica Weith., from Pikermi; Geological Museum, Turin.
; All figures of the natural size.
Fig. b. Side view. Fig. 2. Upper view. Fig. 3. Lower view. Fig. 4. Posterior
upper premolar and molars, right side. Fig. 5. Lower true molars and
posterior premolar, right side.
2. On Two new Genera of Rodents from the Highlands of
Bolivia. By OLprietp Tuomas, F.R.S.
[Received January 28, 1902. |
(Plates VIIT. & IX.)
Mr. Perry O. Simons, the collector who has been doing such
admirable work in the Andean chain, and to whose efforts we
owe the discovery of a very large number of the mammals of that
interesting region, has now sent home a collection from the high
grounds of South-western Bolivia, from the Departments of
Oruro, Potosi, and Sucre. Among these, besides some new species
of known genera, described elsewhere, there are examples of two
rodents so distinct as to demand generic separation, and I have
therefore thought them worthy of being brought before the
Society for description and illustration.
Neocropon, gen. nov. (Octodontide.)
Tail comparatively bushy. Palms and soles granulated, the
pads imperceptible.
Palatal foramina longer than in Octodon, the actual openings,
instead of only the outer fossa, penetrating into the maxille.
Incisors smooth, comparatively thick antero-posteriorly, their
depth rather more than 13 their breadth, as compared with about
1% in Octodon.
Molars rootless, simpler than in Octodon, with a slight concavity
on their outer side and none on the inner, there being no trace
1 For explanation of the Plates, sce p. 117,
LONUINE SBS INVENIO AIN
“HAL SOIg Ute7yULpL “YW 38 Tep FeuG ec
WA Ia 114 4o6l SZ aq
Pe he Se GOW avecoill aie ELSI:
H.Grénvold,del. Photogravure by Bale &Danielsson Lt
1,2,3,44& 64 ANDINOMYS EDAX, ADULT; 47 & 6? YOUNG OF D?
S* & 74 CHINCHILLULA SAHAMA, ADULT; 58 &78 YOUNG OF Dt?
8—I2, NEOCTODON SIMONSI; ADULT.
1902. | NEW BOLIVIAN RODENTS, 115
of the deep internal enamel infolding found in that genus. The
premolar rounded, triangular in section. Lower teeth oval in
section, a slight median constriction on each side of m, and m,,.
This genus is, no doubt, closely allied to Octodon, of which it is
a highland representative.
NEocTopon simonsI, sp. n. (Plates VIII. & IX. figs. 8-12.)
General appearance very much as in the North-American
Neotoma cinerea, strikingly different, by paler colour, whiter
belly, and longer, more bushy tail, from either of the species of
Octodon. Fur soft and fine, hairs of back about 22 mm. in length.
General colour above pale drab-grey, grizzled with black. Sides
clearer drab. Under surface snowy white, well defined laterally,
where it extends rather high up; the bases of the hairs slaty
except on thechin. Front of upper lip hairy, not grooved. Face
coloured like the back, paler on the sides of the muzzle and
cheeks; no definite markings round eyes. A few longer vibrisse
placed just above eye, anda more prominent tuft of them between
eye and ear. Kars large, shaped about as in Octodon, finely
covered with short greyish hairs; a distinct tuft of white hairs at
their anterior bases. Outer sides of arms and legs lke back,
inner sides like belly; upper surface of hands and feet pure
white, the hairs stiff and elongated terminally, so as to surpass
the claws; palms and soles entirely naked, strongly granulated,
with projecting cushions, on which the pads are so small as to be
scarcely perceptible among the granulations; under the heel the
surface is comparatively smooth; pollex with a broad flat nail ;
hallux short, with a claw, its tip falling some way short of the
base of the second digit ; fifth toe, without claw, reaching to the
end of the basal phalanx of the fourth. Scrotum naked. Penis
with a bone, which is flattened, tapering, though not to a point,
about 15 mm. in length. ‘Tail nearly as long as the head and
body, thick, cylindrical, well clothed throughout with hairs, which
increase in length to the end, where they may be fully an inch
long. In colour the tail is brownish black above and at the end,
white proximally on the sides and below; the hairs of the pencil-
end are usually pale brown or even sandy brown, but this appears
to be an effect of bleaching, more or less dependent on the season,
as is the case in the British Squirrel.
Skull and teeth as shown in the figures.
Dimensions of the type, measured by Mr. Simons in the
flesh :—
Head and body 184mm.; tail 152; hind foot, s. u, 36, ¢. u. 38;
ear 32.
Skull—ereatest length 46, basilar length 38, greatest breadth
24; nasals 17 x 5°8; interorbital breadth 10; length of frontal
suture 13°5; breadth of brain-case 18:5; palate length 17:2;
diastema 12; palatal foramina 4°8 x 2:4; length of upper tooth-
series (crowns) 8°3; length of bulla 13; breadth of basti-occipital
on suture 2°6,
g*
116 MR, OLDFIELD THOMAS ON [ Feb, 18,
Hab. Mountainous region south and south-east of the Titicaca-
Poopo basin. Potosi, 4400 metres (type); Oruro, 3700 m. ;
Livichueco, 4500 m.; Challapata, 3800 m.
Type. Adult female. B.M. No, 2.2.2.2. Original number 1620,
Collected October Ist, 1901, by Mr. P. O. Simons. 15 specimens
examined.
Native names ‘“ Chockchuri” and ‘“ Achaco.” ‘ Found among
rocks and cactus, in caves and old Indian tombs; nocturnal”
(Simons).
IT have had great pleasure in connecting with this very beautiful
animal the name of Mr. Simons, in recognition of the remarkable
collecting work he has done in the Andean chain during the last
three years. His collections already number over 1600 mammals,
more than 3000 birds, many hundreds of reptiles and amphibians,
and large numbers of insects and other invertebrates.
ANDINOMYS, gen. nov. (Cricetine.)
Form murine. Thumb with a broad nail. Tail well-haired,
but not pencilled.
Skull rat-like. Muzzle long, broad, and heavy. Interorbital
region narrow, parallel-sided, without ridges, Palatal foramina
large, with very sharply defined edges. Bullee small.
Incisors heavy, smooth anteriorly. Molars very large, highly
hypsodont, as in Chinchillula, but their pattern more as in
Phyllotis, though with almost a microtine appearance in youth,
when they are much more complicated than would be at all
easily perceived from their structure in adult life (see figures).
This genus, like Chinchillula’, is a highly hypsodont and heavily
toothed relative of Phyllotis, itself more hypsodont than the
brachyodont Hligmodontia. But in Chinchillula the teeth are
remarkably simple, practically alike in youth and age, with
opposite and connected enamel-spaces, as shown in the figures
(Plate IX. figs. 5,7). In Andinomys, on the other hand, the
spaces are more or less alternated, the pattern, especially of m,,
becomes less complicated with age owing to the wearing-out of
accessory columns, and the spaces are or gradually become isolated
from each other; the lateral angles are much more acute in
youth, becoming comparatively blunt in old age.
ANDINOMYS EDAX, sp.n. (Plate IX. figs. 1-4, 6.)
General appearance of a large Phyllotis or soft-haired Oryzomys.
Fur long, fine and soft, but ee woolly; hairs of back about 19—
20 mm. in length. General colour above dull buffy or fulvous
buffy, lined oil black; sides clearer, sandy buffy; under surface
not sharply defined, buffy white, the hairs slaty basally. Head like
body ; no orbital markings. ays fairly large, closely haired, brown
1 Chinchillula was originally founded (Ann. Mag. N. H. (7) i. p. 280, 1898) on
a single immature skin, but the British Museum now possesses a series of adult
examples collected by Mr. Simons at Caylloma, Peru.
1902. ] NEW BOLIVIAN RODENTS. 117
outside and in, their edges whitish. Upper surface of hands and
feet well-haired, the hair of the ends of the digits surpassing the
claws, silvery white; fifth hind toe, without claw, reaching to
the middle of the second phalanx of the fourth ; palms and soles
naked, the pads large, rounded and prominent.
Skull with large nasals, very broad anteriorly. Interorbital
region narrow, parallel-sided, concave in the middle line, the
concavity bordered by low rounded and inconspicuous ridges,
which do not overhang the orbit or run back on to the parietals.
Interparietal large and broad. Anterior plate of zygoma-root
concave anteriorly, with an overhanging point above. Palatal
foramina very long and open, broadest mesially, running to a
sharp point behind, where they reach to the level of the first
lamina of m’, their edges very sharp and clearly defined. Posterior
nares level with the back of m’, comparatively broad. Bulle
small.
Dimensions of the type, measured in the flesh by Mr. Simons :—
Head and body 160 mm.; tail 145; hind foot, s. u. 29, c. u. 30;
ear 25.
Skull—greatest length 37; basilar length 31; nasals 15:2 x 6-1;
interorbital breadth (on the convex surface low down) 4, between
the rudimentary ridges 2:1; breadth of brain-case 14; inter-
parietal 4°3 x 10; zygoma-root 3; palate length 18; diastema 10:1;
palatal foramina 9°6 3:2; length of upper molar series 7 ;
combined breadth of upper incisors 3.
Hab. El Cabrado, between Potosiand Sucre, Bolivia. Altitude
3700 metres.
Type. Old female. B.M. No. 2.2.2.15. Original number 1568.
Collected September 20th, 1901, by P.O.Simons. Two specimens.
A young specimen, apparently of the same species, had been
previously obtained by Mr. Simons at La Paz, altitude 4000 m.
Mr. Simons says of this animal: “Caught in thicket of oak-
like bushes; nocturnal.”
Owing to its extreme general resemblance to Phyllotis, the
young specimen from La Paz had been supposed to be an example
of that genus with a wrongly numbered skull, but the later
examples prove that Mr. Simons was in this instance, as usual,
entirely correct in his labelling.
EXPLANATION OF THE PLATES.
Puate VIII.
Neoctodon simonsi, p. 116.
Pruate IX.
Figs. 1, 2, 3. Andinomys edax (p. 116), skull.
Fig. 4. Andinomys edax, right upper tooth-row: a, adult; 4, young.
5. Chinchillula sahame (p.116), right upper tooth-row: a & bas before.
6. Andinomys, right lower tooth-row: a & 6 as before.
_ 7%. Chinchillula, right lower tooth-row : a & b as before.
Figs. 8, 9,10. Neoctodon simonsi (p.115), skull.
Figs. 11, 12. Neoctodow, right upper and right lower tooth-rows.
118 MR. OLDFIELD THOMAS ON NEW [ Feb. 18,
3. On some new Mammals from Northern Nyasaland.
By OupFizLp Tuomas, F.R.S.
[Received February 4, 1902. |
Since my last paper on the mammal-fauna of Nyasaland, a
number of further specimens from that country have been con-
tributed to the National Museum by Mr. Alfred Sharpe, C.B.,
Commissioner, and Col. Manning, Deputy Commissioner, collected
by themselves, Mr. J. McClounie, Mr. J. B. Yule, Capt. Pearce,
and others.
Without occupying space by recording the known species sent,
the present paper, the seventh of the series, gives descriptions of
the new species contained in the collection.
Opportunity has also offered for a re-examination of the
Nyasan Colobus, and, as it proves to be new, it is now described.
CoOLOBUS SHARPEI, Sp. 0.
Coloration exactly as in C. palliatus Peters’, to which this
species has hitherto been referred. Pelage rather closer and
longer, the hairs of the middle back 5-6 inches in length and
those of the mantle 10-12.
Skull decidedly larger in every dimension than that of C. palli-
atus. Frontal region much more convex, and also as it were
higher up the skull, so that the middle pomt between the ridges
is In side view over m” or m°, while in C. palliatus it is over
m; the distance from the same spot to the tip of the nasals is
nearly half as much again in the new form (23 mm. as against 16),
and the nasals themselves are both longer, broader, and less
acutely pointed behind. Zygomata stronger, the vertical height
of the malar just in front of the squamosal suture 10 instead
of about 54 or 6 mm. Zygomatic arches strongly divergent
posteriorly, nearly parallel in C. palliatus. Front edge of
coronoid process of lower jaw angularly convex forwards. The
other cranial differences observable seem all to be dependent on
the greater size of C. sharpet.
Dimensions (approximate) of a stuffed specimen, not the
type :—Head and body 680 mm. ; tail 760; hind foot 190,
Dimensions of the typical skull, that of an old female :—
Greatest length in middle line 115 mm.; condylar length, from
back of condyles to gnathion, 101; zygomatic breadth 85; nasal
opening, height 21, breadth 10; nasals, length in middle line 16,
breadth 10; least interorbital breadth 10; orbit, height 26,
breadth 27 ; vertical height from palate to frontal behind supra-
orbital ridges 38:5; palate length from gnathion 49:5; combined
length of upper premolars and molars 33:6, of upper molars 23,
of lower premolars and molars 38, of lower molars 25.
Hab. Nyasa-Tanganyika Plateau. Z'ype from Fort Hill.
1 Kieured, MB. Ak. Berl. 1879, p. 832, pl. iv 4.
1902.] |. MAMMALS FROM NORTHERN NYASALAND. 119
Type. Old female. B.M. No. 97.7.3.1. Collected by J. B.
Yule, and presented by Alfred Sharpe, Esq. C.B.
It has always been a matter of surprise that the Colobus: of
the high Nyasa-Tanganyika plateau should be the same as that
occurring in the hot lowlands opposite Zanzibar, but the markings
are so nearly identical that no one has hitherto been able to
separate the two forms. Now, however, that the British Museum
has received from Mr. A. B. Percival three fine adult females of
the true C. palliatus from Takaungu, British Kast Africa, with
their skulls, I am able to show that the two are separable, the
skull-differences being really considerable.
I have much pleasure in naming this fine species after my
friend, Mr. Alfred Sharpe, C.B., Commissioner of British Central
Africa, to whose interest and patriotism the National Collection
of Nyasan Mammals is so largely indebted.
Colobus sharpei is the C. angolensis of Sclater (1892), and more
recently the (. palliatus of Pousargues, Neumann, and myself.
HELOGALE VARIA, Sp. nh.
Size rather large. Fur close and fine. General colour above
finely speckled yellowish or buffy grey, passing gradually below
into deep buffy without speckling. Posterior back of all four
specimens, in bleached pelage, dull yellowish or “ clay-colour.”
Head conspicuously different to back, clear deep grey without
yellowish suffusion; a small patch on each side of the muzzle
running back to surround the eye brown or brownish rufous.
Ears grey above, deep yellowish below. Upper surface of hands
and feet dark yellowish clay-colour, scarcely grizzled at all. Tail
coloured like back.
Skull with the nasals broad and parallel-sided for their anterior
half, then abruptly narrowing to a point posteriorly.
Dimensions (approximate) of the type, measured in skin :—
Head and body 270 mm.; tail 162; hind foot (wet) s.u. 46;
ear (wet) 18.
Skull (of the type, nasal sutures still showing) — greatest
length in middle line 53; zygomatic breadth 28°5; nasals 9x 5;
interorbital breadth 9:1; breadth of brain-case above meatus
91:7; palate length from gnathion 24°5; greatest diameter of
p* 52.
: Type. Sub-adult. B.M. No. 2.1.6.5. Four specimens examined.
This Helogale, of which four perfectly similar specimens are in
the collection, differs from all others by the head being much
darker-coloured than the back, these parts being quite concolor in
the other forms. Whether the much greater yellowness of the
rump will also prove.a constant character I cannot say, as in all
the skins the fur of this part is worn and faded.
The recognizable forms of Helogale seem to be as follows :—
(1) HELoGALE ATKINSONI Thos.
_ Helogale atkinsoni. Thos. Ann. Mag. N. H. (6) xx. p. 377 (1897).
120 ON NEW MAMMALS FROM NORTHERN NYASALAND. [ Feb. 18,
_ Face, crown, and back uniformly grizzled grey. Under surface
dull greyish brown, more fulvous on throat and inguinal region.
Feet grizzled greyish proximally, fulvous on digits.
Hab. Abyssinia and Somali.
(2) HELOGALE UNDULATA Peters.
Herpestes undulatus Peters, Reise Mossamb. p. 114, pl. xxv.
(1852).
Body grizzled greyish, more or less suffused with rufous. Face,
especially muzzle, strongly rufous. Crown like back. Under
surface dull greyish brown. Hands and feet deep rufous.
Hab. British East Africa, German East Africa, and Mozam-
bique.
(3) H. vicrorrNa, sp. n.
Body pale grizzled grey, suffused with ochraceous yellow. Muzzle
fulvous. Crown like back. Under surface from chin to anus
dull buffy yellow. Hands and feet also buffy yellow. Tail like
body above, bufty below.
Head and body of type 243 mm.; tail 145; hind foot, s. u. 44.
Skull—ereatest length 52; zygomatic breadth 31; palate length
2:5; greatest diameter of p* 5:5.
Type. Male. B.M. No. 93.5.1.2. Collected October 25, 1892,
by the Rev. F. C. Smith. Presented by Canon Tristram. Three
specimens examined.
Hab. Region of the Victoria Nyanza. Type from Nassa, on
Speke Gulf, south end of the lake. Usambiro, Victoria Nyanza
(Emin Pasha).
(4) H. varia Thos.
As above.
(5) H. parvuna Sund.
Herpestes parvulus Sund. Cifv. K. Vet.-Ak. Forhandl. 1846,
10s LA
Size small. Colour uniformly dark, finely grizzled brown —
throughout, on head, body, belly, and limbs.
Hab. 8.E. Africa: Natal, Zululand, &e.
FUNISCIURUS YULEI, sp. n.
No stripes. Premolars 2.
Superficial appearance somewhat as in /’. cepapi. Size rather
larger than in that species. Fur of medium harshness; hairs of
back about 10 mm. in length. General colour above pale coarsely
grizzled tawny, greyer over the shoulders, though this may be
due to old age. Sides greyish tawny, much lighter than the
dorsal area, though the line of distinction is not sharply defined.
Under surface ill-defined whitish, whiter on the chest and groins,
mixed with greyish on the belly. Sides of muzzle and rings
round eyes whitish. Crown grizzled tawny. Ears conparatively
US DEVE els LNG alah Ib Sher eiilval ss WOlel 721
‘dor sozg uses THERE G EE Ee) apstertss Ip ar
Kid’ 1194" 2061 S Za
“SWINCIWel el —SVOIMIN/ SEINE S) Gl “IESE WGINEE Ele “SAL DIFDINOO) SinarEtuiet 22 WOvEl Ih
‘Gar sozgurequyy, ’ UEEFE EID speess) Pal
TEC Wel TES COGIL SZ L
; 1902.] ON THE YOUNG OF POLYPTERUS. 121
large, whitish, especially at their edges. Armsand legs like sides ;
hands and feet heavily built, their upper surfaces greyish white ;
soles naked except under the heels. Tail but little bushy, its
hairs broadly ringed with black and pale yellowish, their tips
whitish.
Skull rather heavier than that of F. cepapi. Nasals broad,
expanded posteriorly. Interorbital region broad, flat or slightly
concave ; postorbital processes well developed, projecting directly
outwards instead of backwards. Premolars 2. Molars heavier
than in J’. cepapi.
Dimensions (approximate) of the type, measured in skin :—
Head and body 205 mm.; tail 145; hind foot s. u. (wet) 41; ear
(wet) 19.
Skull—henselion to basilar suture 28; nasals, length 12:5,
least breadth 5, posterior breadth 6; interorbital breadth 12;
tip to tip of postorbital processes 19; intertemporal breadth 13 ;
palate length from henselion 18°5; diastema (to front of p') 10:6;
length of tooth-row (omitting the small p’) 8. Lower jaw, condyle
to incisor-tip 27:7.
Hab. Muezo, near Lake Mweru.
Type. Old male. B.M. No. 2.1.6.8.
It is difficult to say to which species /. yulet is really most
closely allied. Externally it has a certain resemblance to F’. annu-
latus, but that animal has only one upper premolar and differs in
many other details. From /. cepapi, found in the same region,
it is distinguished by its larger ears, grey instead of fulvous
limbs, whitish feet, larger molars, and other points both external
and_cranial.
I have connected with this distinct Squirrel the name of
Mr. J. B. Yule, of the official staff of the Protectorate, by whom
a large number of the North Nyasa specimens described now and
in former papers have been collected.
4, Qn some Characters distinguishing the Young of various
Species of Polypterus. By G. A. Boutzncsr, F.R.S.
[Received January 28, 1902. |
(Plates X. & XI.)
The increased interest which has lately been paid to the
remarkable African Crossopterygian Polypterus has resulted in a
better understanding of the characters by which the species can
be distinguished *, and the recent exploration of the Congo has
added several forms, fully entitled to specific rank, which were
undescribed °,
1 Wor explanation of the Plates, see p. 125.
2 Cf. Boulenger, Ann. & Mag. N. H. (7) 11. 1898, p. 416.
3 af Boulenger, Poiss. du Bass. du Congo (1901), and Ann. Mus. Congo, Zool. ii.
(1902).
122 MR. G. A. BOULENGER ON THE [ Feb. 18,
We can now distinguish ten Polyptert :—
1. P. bichir Geoffr.—Nile. ;
2. P. lapradi Stdr.—Senegal, Gambia, Niger.
3. P. congicus Blgr.—Congo, L. Tanganyika.
4, P. endlicheri Heck.—Nile, Niger.
5. P. delhext Blgr.—Congo.
6. P. ornatipinnis Blgr.—Congo.
7. P. weeksii Blgr.—Congo.
8. P. senegalus Cuv.—Nile, L. Rudolf, Senegal, Gambia,
Niger.
9. P. palmas Ayres.—West Africa, from Liberia to the
Congo.
10. P. retropinnis Vaill—Congo.
Not before 1869 was anything known of the characters of the
young. We owe the first observations on this subject to Stein-
dachner !, who, on his return from a collecting expedition to the
Senegal, announced the startling discovery that both P. lapraduw
and P. senegalus are provided, for a certain period, with a large
external opercular gill similar in structure to those possessed by
Tailed Batrachians. In 1896 °*, I noticed the presence of external
gills in P. palmas, and successively in specimens of P. congicus,
lapradu, and weeksii*. In his highly interesting memoir on the
Breeding-habits of some West-African Fishes“, Mr. Budgett has
made us acquainted with the external appearance of a specimen,
referred by him to P. lapradii (but which he informs me, after
examination of the Nigeria specimens sent by Dr. Ansorge,
should be referred to P. senegalus), smaller than any previously
obtained and which may be regarded as truly larval. Having
recently received, from various sources, a number of young
specimens from the Nile, the Congo, and Nigeria, I am able to
supplement our present knowledge on various points. The notes
here offered deal with six species: P. lapradu, P. congicus,
P. endlicheri, P. weeksti, P. senegalus, and P. palmas. It will be
observed that the young of P. bichir, the oldest known species,
the only one to occur in the Lower Nile, is still undescribed.
Before proceeding to descriptive details, I wish to observe that
the specimens with external gills at present known may be
arranged in three divisions:—(1) Without scales, and with the
dorsal fin spineless, not differentiated from the caudal; the only
example known being the larval specimen of P. senegalus, brought
home from the Gambia by Mr. Budgett and described and figured
by him. (2) With scales of a cycloid type and with the dorsal
fin as in the preceding. (3) With all the essential characters
of the mature form. Although the scales may, in the very
1 Sitzb. Akad. Wien, lix. i. 1869, p. 103.
2 Ann. & Mag. N. H. (6) xvii. 1896, p. 310.
3 Ann. & Mag. N. H. (7) ii. 1898, p. 419, and P. Z. 8S. 1898, p. 493, 1899, p. 554,
1900, p. 267.
4 Trans. Zool. Soc. xvi. 1901, p. 116.
1902.] YOUNG OF POLYPTERUS. 123
young, differ so greatly in shape, their numbers are the same
as in the adult, and, though devoid of spines and in no. way
“pinnules,” the rays of the dorsal fin are identical in number ;
only, as it is practically impossible to establish a limit between
them and those of the caudal, it is preferable to count the rays
right to the extremity of the vertebral column.
POLYPTERUS LAPRADII Stdr. (Plate X. figs. 1 & 2.)
This is one of the largest species, growing to a length of
740 millim. The largest specimen with fully developed opercular
gills, obtained by the late P. Delhez at Kaédi, Senegal, measures
300 millim., and is the largest Polypterws with external gills yet
recorded ; the specimens in which the external gills were discovered
by Steindachner measured up to 230 millim. These gills are
retained, as a rule, until the young is 240 to 260 millim. in
length, but they vary in the degree of development irrespective
of the size of the specimen, and sometimes also on the two sides.
Four young specimens, measuring 94, 98, 114, and 205 millim.
respectively, were obtained at Assay and Abo, Southern Niger la,
in October last by Dr. W. J. Ansorge, to whom ichthyology is
indebted for so many striking discoveries in that part of Africa.
In the smallest specimen the external gill measures 30 millim.,
the dorsal rays are all simple and spineless, 23 in number, and the
scales are very thin, circular, juxtaposed, and only well developed
about the lateral line and on the tail; the caudal fin is acutely
pointed, the median rays being produced and as long as the head.
A black band extends on each side from the end of the snout,
through the eye, to the base of the external gill, which is likewise
black, and along the body to the base of the caudal fin ; this band,
on the body, is about as broad as the eye; below it a narrower
black band extends from the shoulder to the anal fin. In the
two next specimens the external gills measure 33 and 50 millim.
respectively. In the largest specimen, the right external gill
measures 68 millim., the left 53; the scales are well developed,
rhomboidal and imbricate, and the dorsal spines (14 in number,
followed by 10 soft rays to the extremity of the vertebral column)
are ossified and bicuspid, supporting three articulated rays to
form the “pinnule.” The dark bands are more indistinct and
crossed by bars on the caudal region.
PoLyPTERUS concicus Blgr. (Plate XI. fig. 1.)
This species appears to be the largest of the genus, growing to
one metre. I have already reported upon specimens up to 260
millim. provided with the external gill, and one of them has
been figured in the ‘Annales du Musée du Congo.’ It appears
that in this species, as in P. lapradwi, the external gills are
normally retained until comparatively late. A specimen recently
received trom Banzyville on the Ubangi, and belonging to the
Congo Museum, is interesting as veer the smallest yet obtained
of that species.
124 MR. G. A. BOULENGER ON THE [ Feb. 18,
It measures 118 millim., the external gill being 25 millim.
long. In its development it is intermediate between the two
stages noticed above in P. lapradii. The dorsal spines are not
yet defined, the number of rays being 22 to the extremity of
the vertebral column; the scales are imbricate, cycloid, with a
tendency to the rhomboid shape, rugose, with a smooth central
area corresponding to the part of the scale first to appear. Six
blackish bars across the back, bifurcating on the sides; below
these bars, two interrupted blackish lines run along each side.
PoOLYPTERUS ENDLICHERI Heck. (Plate XI. fig. 2.)
I have seen only one young specimen of this Polypterus,
obtained at Abo, Nigeria, by Dr. Ansorge in October. It
measures 180 millim., the external gill 30, and corresponds in its
development with the largest specimen of P, lapradi obtained by
Dr. Ansorge in the same locality. The spines number 11, and
are followed by 8 soft rays. The coloration does not differ from
that of the adult.
PoLYPTERUS WEEKSI Bler. (Plate X. fig. 3.)
The type on which this species was established in 1898
is a young specimen, 170 millim. long, with external gills
measuring 15 millim., obtained at Monsembe, Upper Congo,
by the Rev. J. H. Weeks, from whom the British Museum has
since received a specimen 380 millim. long and without external
gills, obtained at the same locality. The young specimen, which
is here figured, is in what I have defined above as the third
period. It is dark olive above, yellow beneath, the two colours
sharply delimited on the side; six narrow black bars across the
back, with some black spots between them, the last followed by
irregular marblings; a large black spot on the membrane to each
dorsal spine; soft fins with dark and light spots; pectoral with
three blackish cross-bands. I have already observed that the
asperities on the scales, which are so striking in the young,
disappear in the adult.
PoLYPTERUS SENEGALUS Cuv. (Plate XI. fig. 3.)
This species differs from all its congeners in being of a uniform
greyish olive, without any markings, at least in specimens above
120 millim. total length; but the very young, with which we
have only quite recently become acquainted, are striped, as
described hereafter. The external gills appear to be lost very
early ; for they have not yet been found in any specimen above
90 millim., although a large number of young have been examined
by Steindachner, Budgett, and myself.
Two small specimens, measuring 60and 69 millim. respectively,
were obtained by Mr. Loat in the White Nile, at the mouth of
Lake No, in the beginning of February 1901. In both, the
external gills are present and measure 5 millim. The smaller
specimen has 8 spines and 8 soft rays to the dorsal, the larger
1902.] YOUNG OF POLYPTERUS, 125
9 spines and 7 soft rays; in both the 5 anterior spines are not
fully formed, not bicuspid, whilst the others have already assumed
their definite shape; the caudal fin is long, with the median rays
produced, a little longer than the head; the scales are imbricate
and rhomboidal, striated, except on the central areola. A dark
brown band on each side, from the end of the snout, through the
eye and over the external gill, to the base of the caudal; another
dark band, only a little narrower, below the first along the body,
separated from it by a narrow yellowish streak. The very young
of P. senegalus is therefore banded like that of P. lapradii, with
this difference, that the lower band is broader in proportion to
the upper.
Dr. Ansorge’s Nigeria collection contains several young with
external gills, obtained at Abo in October 1901; the length of
these specimens varies between 39 and 105 millim. In the
largest the external gill is present only on the left side and
measures 9 millim.; the dorsal spines, 9 in number, are bicuspid,
the scales are nearly smooth, and mere traces of the dark bands
are visible: the other specimens agree entirely in their markings
with those obtained by Mr. Loat in the White Nile, and show
the complete passage from cycloidal to rhomboidal scales, and of
simple dorsal rays to pinnules with spines, the posterior of the
latter being developed first.
PoLyPTERUS PALMAS Ayres. (Plate XI. fig. 4.)
This species is closely allied to P. senegalus, and in it also the
external gills do not persist long. I have only observed them in
one specimen 95 millim. long, where they measure 8 or 9 millim. ;
in all other specimens examined, measuring 80 millim. and above,
I have found them to be absent.
The body is brownish above, yellowish beneath; the upper
parts are marked with darkex cross-bars, close together, which
branch off into marblings or «, wide-meshed network on the sides,
enclosing roundish yellowish spots; or the dark and light spots may
be arranged in chess-board pattern on the sides; a blackish oval
spot on the peduncle of the pectoral fin.
EXPLANATION OF THE PLATES.
PuatTE X,
Fig.1. Polypterus lapradii Stdr., p. 123, young from Abo, S. Nigeria, slightly
reduced.
2. Younger specimen of the same species, from Assay, S. Nigeria, nat. size, with
(2 a) enlarged view of scales from the middle of the side.
3. Polypterus weeksi Bler., p. 124, young from Monsembe, Congo, nat. size,
with (3 a) enlarged view of scales from the middle of the side. :
Prater XI.
. Polypterus congicus Blgy., p.123, young, from Bangyville, Ubangi.
. Polypterus endlicheri Heck., p. 124, young, from Abo, S. Nigeria.
. Polypterus senegalus Cuv., p. 124, young, from Assay, S. Nigeria.
. Polypterus palmas Ayres, p. 125, young, from Monsembe, Congo.
[All figures of the natural size. |
Poo lO
126 . ON A NEW AFRICAN SNAKE. [Feb. 18,
5. Description of a New Snake of the Genus Psammophis,
from Cape Colony. By G. A. BounEnexr, F.R.S.
[Received February 17, 1902. ]
(Plate XII.)
PsAMMOPHIS LEIGHTONI. (Plate XII.)
Snout once and two thirds as long as the eye, with a shallow
concavity in front of the vertex. Rostral a little broader than
deep, visible from above; nostril between three shields; inter-
nasals shorter than the prefrontals; frontal twice as long as
broad, in the middle about two thirds the width of the supra-
ocular, as.long as its distance from the end of the snout, slightly
shorter than the parietals; loreal twice as long as deep; a single
preocular, forming an extensive suture with the frontal; two
postoculars; temporals 2+2; eight upper labials, third deeper
than second and fourth, fourth and fifth entering the eye; four
lower labials in contact with the anterior chin-shields, which are
shorter than the posterior. Scales in 17 rows. Ventrals 156;
anal divided; subcaudals 84. Dark brown above; the middle
row of scales black with yellow shafts forming an interrupted
light vertebral line; a yellow lateral streak along the adjacent
halves of the third and fourth rows of scales; the upper half of
the fourth scale black; scales of outer row yellow in front and
brown or black behind; sides of neck with dark ocelli edged
with bright yellow; head dark brown above, with a yellow line
along the middle of the snout and another on each side of the
frontal shield; two pairs of yellow spots on the parietal shields ;
four yellow bars on each side of the head, the first on the pree- °
ocular, the second on the postoculars, the third extending to the
upper surface of the head and nearly meeting its fellow on the
occiput; rostral and labials yellow, with black spots; lower parts
yellowish white, with black dots and two bluish-grey longitudinal
lines which widen forwards into two bands and unite on the
throat.
Total length 910 mm. ; tail 270.
A single specimen, from Eerste River Station, 21 miles east
of Cape Town, was received by Dr. G. Leighton, of Pontrilas,
Hereford, and presented by him to the British Museum.
Tn its markings this Snake differs from all its congeners. It
differs from P. sibilans in having the rostral shield broader than
deep, as in P. notostictus, in which the preocular and the anal
are divided.
EXPLANATION OF PLATE XII.
Psammophis leightoni. Upper, lower, and side views of head and anterior part of
body and upper view of middle part of body. Natural size.
f
PLZ. 02 soll 1b sel 2 .
tern Bros .1mp.
VG
PJ. Smit del.et hth.
PSAMMOPHIS LEIGHTONI.
1902.] ON CARPAL VIBRISSH IN MAMMALS. 127
6. Observations upon the Carpal Vibrisse in Mammals. By
Frank EK. Bepparp, M.A., F.R.8., Vice-Secretary and
Prosector of the Society.
[ Received December 24, 1901. ]
(Text-figures 17—21.)
In a brief note in ‘Nature’', and incidentally in a paper
’ yy pap
devoted to the anatomy of Bassaricyon’, I directed the attention
of zoologists to a tuft of long and strong hairs which exist in
many mammals on the wrist close to the root of the thumb and
generally on that (the radial) side of the forearm. These long
hairs are quite similar in character to those which are found in
various parts of the head and face of many mammals, such as, for
example, the “‘ whiskers” of the domestic cat. They are readily
seen on account of their size; and, as a rule, they are also
conspicuous by reason of the fact that they are frequently, though
not always, of a different colour from the hairs of the surrounding
pelage. Butif they escape the eye, as is sometimes the case in the
skins of spirit-preserved specimens, they can be felt through the skin
on account of the large hair-bulbs which receive the proximal ends
of the hairs. That these structures must be of some use to their
possessors seems to be obvious, and yet is not easy to prove. I
have watched various animals, and cannot see that they make any
use of the tuft of hairs upon the wrist for touching objects,
except in the possible case of the Raccoon (Procyon lotor), which
did appear to me to hold its food rather nearer to the wrist than
is usual with animals. I believe that my two brief notes referred
to are the first published statement of the general presence of this
carpal tuft of hairs in mammals. Some years since, as I have
already acknowledged, Mi. Bland Sutton described these hairs in
various Lemurs*, and showed plainly that they are a general
character of that group, though they were wanting (and I can
here confirm Mr. Sutton) in the Potto, I find, however, that in
every group of animals, with the exception of the Apes, which
use their front limbs as grasping-organs as well as for locomotive
purposes, these structures are present with some few, though rather
striking, exceptions.
Since the publication of the facts contained in those two papers,
I have had the opportunity of examining a large number of
mammals belonging to various Orders. I am therefore now in a
position to extend the statements which I originally made, and
to give more in detail the distribution of these curious structures
in the group of mammals. I do not think that we have here a
secondary sexual character, though it is possible that in some
1 Vol. lxii. p. 523.
2 “On the Anatomy of Bassaricyon,” P. Z.S. 1899, p. 661.
3“On the Arm-Gland of the Lemurs,” P. Z.S. 1887, p. 869; where they are
figured in the genera Hapalemur, Lemur, and Cheirogaleus,
128 MR, F, E. BEDDARD ON [Feb, 18,
cases the male may be provided with the organ which is wanting
in the female. I called the attention of the Society some little
time since to the fact that the female of Hapalemur simus* has
not this tuft, which is very plain and obvious in most Lemurs,
Text-fig. 17.
Left fore foot of Dasypus villosus; ventral surface.
H, tutt of vibrissz.
including the male of the closely allied H. griseus. I have,
however, met with the structure in too many females of different
species of mammals to allow of its being regarded as a sexual
1 “ Notes on the Broad-nosed Lemur, Hapalemur simus,” P. Z,S, 1901, i. p. 121,
1902. ] CARPAL VIBRISS4 IN MAMMALS. 129
character. Broadly speaking, it may be stated that this sense-
organ, aS we may in the meantime assume it to be, characterizes
the Lemurs, Rodents, Carnivora, and Mayrsupials, and that it is
absent in the Ungulates (with the exception of Hyraa:) and in
the Primates (excluding the Lemurs), The Bats I have not yet
studied from this point of view. Of the Insectivora I am un-
willing to speak, as I have examined only Cenietes and Lrinaceus,
which certainly had not this tuft of hairs.
As to the Edentata, the representative of this carpal tuft of
vibrisse does appear to exist at least in the Armadillo, Dasypus
villosus, as the accompanying figure shows (text-fig. 17, p. 128).
But the hairs of that mammal are so coarse that there is but little
difference in size and general appearance between the tuft which
I compare to those of such an animal as Petaurus sciureus (text-
fig. 18, p. 130) and the general hairs of the body. The Sloth
(Bradypus tridactylus) has not any traces that I could discover of
this “organ.” As to other Edentates I have no information to
offer, except concerning Manis, where I have found no traces of
these hairs.
Considered broadly, therefore, this carpal tuft of vibrissz is of
some little use for classificatory purposes, apart from its absence in
the Ungulates, where it might well be supposed to be deficient on
account of the lack of facilities for use. The most salient feature
as to its absence or presence is its nearly universal existence
in the Lemurs, and the absolutely universal absence (as far as I
have ascertained) in the Monkeys. These two divisions of the
Primates, as they are most commonly considered to be, have
been brought nearer to each other by recent researches upon
certain extinct forms such as Vesopithecus*, and by investigations
upon the placenta of Zarsiws, which has been shown to be ape-
like and even human in its characters’. It is not, therefore,
without interest to be able to bring forward a character which
seems to absolutely distinguish these two divisions of the Pri-
mates. Furthermore, it is not a character which has an obvious
relation to ways of life: if the tuft of vibrissie is useful to the
Lemurs, it would seem to be equally useful to the Monkeys, many
of whom use their hands as climbing- and grasping-organs in the
same way. And I can at least assert, that while the majority of ©
the Lemurs (excluding the Potto and the Loris) which I have
examined possess this tuft, the large number of Monkeys, both
of the Old and New Worlds, which have passed through my
hands do not possess it.
As to the Marsupials, the genus J/acropus, so far as my present
investigations go, stands alone in that the wrist is not provided
with this tuft of vibrisse. I have examined both adults and
quite newly born individuals of several species. In the naked
new-born young of Marsupials this tuft of vibrisse is exceedingly
1 Worsyth Major, P. Z.S. 1899, p. 987.
2 See, for a survey of the position of Tarsius, Harle, Amer. Naturalist, xxxi. p. 569.
Proc. Zoon. Soc.—1902, Vou. I. No. IX. 9
130 MR. F, E. BEDDARD ON [ Feb. 18,
obvious, and is nearly the only vestige of hair visible to the naked
eye that the very young Zpiprymnus, Potorous, and Phalangista
possess. But this carpal sense-organ is by no means confined to
the Diprotodontia. I have found it in Perameles and im the
Rat-tailed Opossum (Didelphys nudicaudata).
The Carnivora as a group are apparently characterized by the
existence of this organ. I have found it in representatives of
the Ailuroidea (Domestic Cat, Lion, Cynictis levaillant, Herpestes
Text-fig. 18.
Right fore foot of Petawrus sciureus; ventral surface.
H, tuft of vibrissz.
pulverulenta), the Arctoidea (Putorius, Otter, Mustela, the Coati),
but I do not find it in the Dogs. There are, however, apart from
the Dogs, some exceptions to its occurrence in the Carnivora. It
certainly does not exist either in Viverra civetta or in V. tanga-
lunga. Oddly enough, I could not detect the tuft of hairs in the
Tiger, obvious though they are in the Lion.
The Rodents again are, as a rule, to be characterized by the
1902. | GARPAL VIBRISSA IN MAMMALS. 131
possession of this tuft of vibrisse. Its presence is not, however,
universal in this order of mammals. On the whole, I should be
disposed to think that those Rodents whose feet have more of an
Ungulate character, such as the Capybara and Dolichotis, are
without the structure in question; but tbe majority of the genera
which I have examined certainly are furnished with these long
vibrissee. I have found them, for example, in several species of
Sciurus, in Cricetus, Gerbillus, Microtus, Saccostomus, Acomys,
Mus, and Pteromys. It is remarkable that though they exist in
Text-fig. 19. ;
Right hind foot of Petawrus seiwreus; lateral surface.
H, tuft of vibrisse.
the Flying-Squirrel just mentioned, and I have had the oppor-
tunity of verifying their occurrence in two species, they are
totally absent from the wrist of another genus of Flying-Squirrel,
viz. Sciwropterus. Nor can I find the tuft in the Jerboa or in
Rhizomys.
Another peculiarity of this tuft of specialized hairs is that it is
not invariably to be found in all individuals of a given animal.
As a rule, so far as my experience goes, it is the case that the tuft
is constant and to both sexes. But in the Suricate I have found
132 MR. F. E. BEDDARD ON [Feb. 18,
the tuft absent in one example, a male, and present in a » second
of the sex of which I have no record.
Tn all the cases that have been referred to in the above brief
Text-fie. 20.
Left tore foot of Nasua narica; ventral surtace.
Dissected to show nerve (N) supplying tuft of vibrisse (H).
1902. | CARPAL VIBRISS@ IN MAMMALS. 133
summary of the facts at my disposal, the vibrissze were only to be
found in the fore limbs. In some cases I admit I did not ex-
amine with great care the tarsus as well as the carpus. But after
finding that in one species (illustrated in text-figs. 18, 19, pp. 130,
131) both fore and hind limbs showed a precisely similar tuft of
vibrisse, I naturally examined other animals that came my way.
In Petaurus sciureus, in fact, this tuft of sensory hairs is present
on both pairs of limbs, occupying a corresponding position in each.
There was no difference that I could detect in the arrangement
or structure of the vibrissee in the two limbs. The case appears
to me to be simply another instance of structures appearing in
one pair of limbs being repeated in the other,—just as the horny
spine-like outgrowths of the wrist of apalemur griseus are found
on the ankle of Galago garnetti*.
The accompanying illustration (text-fig. 20, p. 132) shows the tuft
of carpal vibrissee in a Coati with the skin-flap removed and the
muscles partly dissected. In this animal the tuft consists of six
long black hairs, the implantation of which on the skin is shown
in the drawing. The roots of these vibrisse are provided with a
nervous supply i in the shape of a strong branch leaving the main
nerve of the arm just opposite to the tuft, and ending after a short
course without any sensible diminution of thickness and without
any branching visible to the naked eye. This marked nervous
supply to the carpal sense-organ is not, however, invariably found.
The most aberrant condition of this carpal organ in all the animals
which I have examined exists in Hyrax. In this Ungulate, the
only Ungulate in which I have been able to detect the or gan at
all, I could only find it on one of the two fore limbs ; and on that
limb, the right, it was represented by only two long hairs, one
being situated in the normal position which the tuft occupies in
other mammals, and the other placed some way behind this. On
carefully dissecting away the skin the roots of the hairs were ex-
posed, and their great size enabled it to be seen that there were
two of them, though only one hair was visible externally. In the
case of the posterior tuft I could find but one han-bulb. This
state of affairs is shown in the drawing exhibited herewith (text-
fig. 21, p. 134). The most careful examination fa‘led to show
any nerve-branch supplying the roots of the vibrissee. I have no
doubt that minute microscopic threads exist ; but there is nothing
that can be detected with the naked eye to be seen; and I think
that it could hardly have been overlooked. Now it appears to
me that we have in this animal a case of commencing retro-
gression of the organ in question. It can hardly be denied that
Hyrax stands at least nearer to the base of the Ungulate series
than do the Perissodactyla or the Artiodactyla. Therefore it is
not surprising to find in Hyrax traces of a structure that has
entirely disappeared in the more specialized forms; and in effect
1 See Beddard, P. Z.S. 1884, p. 393, and ibid. 1901, vol. i. p. 2
134 MR. F. E, BEDDARD ON [ Feb. 18,
this appears to me to be the case. In Hyraa there is still a
considerable trace of the carpal sense-organ, present (?) in the
ancestral Ungulates, which had not completely left off the use of
Text-fig. 21.
Left fore foot of Hyrax ; ventral surface.
H anterior, H’ posterior vibrissx.
their fore limbs as grasping-organs; but it is evidently under-
going degeneration, and has finally disappeared in the newer forms
of Ungulates. At least it has disappeared, so far as my
1902. | CARPAL VIBRISSE# IN MAMMALS. 135
observations go, in the form which it jossanse? in the Carnivora,
Marsupials, &e.
But I shall now proceed to urge some facts and suggestions
which would tend to show that in the Horse tribe there are traces
of this organ present in the well-known “chestnuts” of those
animals—callosities on the fore and hind feet or fore feet only.
And to do so I must revert to the Armadillo. The condition of
this organ in the Armadillo requires some further description, as
it differs in certain points from what is to be found in other
mammals. As will be seen from the drawing (text-fig. 17, p. 128),
the long vibrisse are not so markedly longer than the hairs which
clothe the skin generally as is the case with other mammals. The
general hairy covering of the Armadillo is coarse. In the second
place, they are decidedly more numerous and not arranged in a
tuft; they do not, that is to say, apparently spring from the same
circumscribed spot. On the contrary, they are borne upon a
raised patch of integument which is about half an inch long;
this tract of integument, moreover, is considerably thickened,
which marks it off from the surrounding integument in a very
distinct way. A dissection of the skin in this region shows a
nerve supplying this tract of vibrissee-covered skin; but the nerve
is rather small in proportion to its bulk in such an animal as
the Coati (text-fig. 20, p. 132). The tract of skin bearing the
vibrissee would be quite obvious if it bore no vibrisse at all. It
has, too, a hard “feel.” Now if this specialization of the pad
bearing the vibrissze were to proceed further, it would become a
mere horny pad and the vibrissee would cease to grow upon its
general surface, as with the pads of the foot. They would, so to
speak, be driven off or at least to one corner. The resulting state
of affairs would be such as is represented in the Lemur, where
(Lemur catta)there is a horny pad, to the side of which is the
tuft of carpal vibrisse; a still further specialization would of
course bring about the conditions which I and Mr. Sutton have
shown to obtain in Hapalemur griseus. On the other hand, the
disappearance of the tuft of vibrissee would result in a structure
precisely like the ‘‘chestnut” on the fore limbs of the Equide.
It appears to me that the condition of the sensory organ upon
the wrist of Dasypus villosus distinctly suggests a possible origin
for the “chestnuts” of the Horse tribe, which have been variously
explained, but not in this way. There is every @ priori likelihood
of finding traces in the Ungulata of this widely spread organ, and,
as already stated, in the primitive ungulate Hyraa they actually
exist, in a but slightly modified form, The difficulty caused by
the fact that the hind limbs in the true Horses have also “ chest-
nuts ” is removed by the occurrence in Petawrus sciureus of these
organs on the hind as well as on the fore lambs’.
1 Jt must be borne in mind, however, that in the horse the “chestnuts” of the
hind limb are upon the ankle, those of the fore limb above the wrist. The Jatter
position is that of other mammals.
136 ON CARPAL VIBRISS® IN MAMMALS. [ Feb. 18,
It may be convenient to sum up the facts which have been
detailed in this communication, and to present them in the form
of a brief réswmé :—
(1) In nearly all the Orders of Mammalia—viz., the Primates,
Carnivora, Ungulata, Rodentia, Edentata, and Marsupi-
alia—there is generally a tuft of strong vibrissee upon the
wrist.
(2) This tuft consists of from one to about twenty hairs usually
(if not always) supplied by a strong nerve which arises
from the radial nerve of the arm.
3) This structure is as a rule, when present, found in both
) p )
sexes ; but occasionally it is present in the male only.
(4) The tuft of carpal vibrissee is apparently absent from all the
Ungulates, except Hyraz; and from the Anthropoidea
among the Primates.
(5) In the groups where the carpal organ is present, 1t is apt
to be capricious in its distribution. Thus it is present in
the Lion, and absent in the Tiger.
(6) With the marked exception of the Anthropoidea, there
seems to be a certain relation between the presence of
the tuft of carpal vibrissee and the nature of the fore
limbs. When the fore limbs are purely ambulatory limbs,
the carpal vibrisse are absent.
1902.] THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 137
March 4, 1902.
W. Bateson, Esq., F.R.S., Vice-President, in the Chair.
The Secretary read the following report on the additions to the
Society’s Menagerie during the month of February 1902:
The registered additions to the Society’s Menagerie during the
month of February 1902 were 73 in number. Of these 24 were
acquired by presentation and 49 were received on deposit. The
total number of departures during the same period, by death and
removals, was 154.
Amongst the additions attention may be specially directed to :—
(1) A young male Snow-Leopard (felis wneia), from Ladakh,
presented by Capt. H. I. Nicholl, of the 1st Bedfordshire Regiment,
Mooltan. We are much indebted to Capt. Nicholl for the care
and trouble which he has taken in sending us home this rare and
beautiful animal, of which so few specimens have ever reached
us, and to Messrs. Gellatly, Hankey, & Co. for its passage home in
the s.s. ‘Prome.’ It was originally obtained by Capt. Nicholl
in Ladakh about September 19011.
(2) A pair of Prjevalsky’s Wild Horses (Lquus prjevalskit),
being part of the same convoy as those lately acquired by His
Grace The Duke of Bedford, our President, as was recently
announced (see P. Z.S. 1901, vol. ii. p. 505).
Mr. Hagenbeck has supplied me with the following information
respecting the capture of these animals :—
The Wild Horses were captured in three different districts in
the vicinity of Kobdo in Western Mongolia (in about 38° N. and
90° 35’ E.), in the Chinese Empire, as shown on the map which I
now send.
From Kobdo the horses were taken to the Siberian Railway
Station, Ob. They were thirty-nine days on the way, including
four days of travel by barges on the River Ob.
The Prjevalsky’s Horses drop their young ones from the latter
days of April to about the 20th of May, and during this time
they come to particular spots, which are marked on the map, and
they find at these places plenty of food and water.
The system of catching them is the following :—Large troops
of Mongols hunt in combination, waiting for the animals behind
the hills, and when they observe that a great many are together
‘they all, on a signal, suddenly start and ride after the animals.
As the young ones cannot follow their mothers, they are caught
with nooses that are arranged on long sticks. The captives are
brought to the camp, where the Mongols keep a lot of common
mares with their young ones. These young ones are then taken
away, and the wild colts put to the common Mongol horses to
be nursed by them. After a few days the young ones become
1 See ‘The Field’ of 1902 (vol. xcix. p. 825) for an account of the capture of this
animal.
Proc. Zoot, Soc.—1902, Vou, I. No, X, 10
138 MR, G. T, BETHUNE-BAKER ON [ Mar. 4,
well acquainted with their nurses and then follow easily up to
Kobdo.
I exhibit a water-colour drawing (Plate XIII.) by Mr. Smit,
taken from two of the specimens now in the Gardens, which gives
a good idea of the general appearance of Prjevalsky’s Horse in its
winter dress.
Mr. B. Tegetmeier, F.Z.S., exhibited a series of photographs of
Prjevalsky’s Horse, taken on different occasions, and stated that
further information on the subject would be found in ‘ The Field’
of January 11th (vol. xcix. p. 69, 1902).
Mr. E. N. Buxton, F.Z.S., exhibited a series of photographic slides
illustrative of bird- and animal-life on the White Nile, which he
had lately visited. He called attention to the enormous abundance
of aquatic birds and waders which resort to the mud-banks of the
Nile south of Khartoum and to the numerous lagoons in the
marshes in the neighbourhood of Fashoda. Portraits were shown
of Pelicans, Sacred Ibises (worshipped by the Egyptians of the
early dynasties, but not now found in Lower Egypt), and many
other birds—such as White Ibises, Black Ibises, Buff-backed
Herons, Jabirus, Cranes, Stilts, Fish-Eagles, Goliath Herons, and
Marabous. These had been secured by the use of a telephone-
lens (by Dallmeyer), as also portraits of the Water-buck, the
White-eared Cob (Cobus leucotis), the Tiang (Damaliseus tiang),
and the Hippopotamus. ‘The difficulties of making an approach
to these wild animals for photographic purposes were described.
Some characteristics of the Roan Antelope (Hippotragus leuco-
pheus) were pointed out, and the habits of the Buffalo of the
Nile and of the Reed-buck (Cervicapra bohor) were explained.
Mr. Buxton expressed the opinion that the habit of the natives
of burning the grass on the marshes affected the coloration of the
White-eared Cob. The variation in colour of individuals of this
species was illustrated by two heads of old males, and a skin of
another individual was exhibited showing a white spot on the
withers resembling that found in Mrs. Gray’s Water-buck (Cobus
Maria).
Denes were also shown of village-life among the Dinkas—a
race of remarkable stature which inhabit the southern part of the
Ghezireh. .
Mr. G. T. Bethune-Baker, F.Z.8., presented a paper, entitled
“A Revision of the Amblypodian Group of the Butterflies of
the Family Lyceenide,” and made the following remarks :—
This important section of the Lyczenide is broadly confined to
the Indo-Malayan region, though a few species are found in
China, Japan, and Queensland, and a few have also been recorded
WAS A ANC Siete CS ililie) eat
{ Lucy SOME UAO QU TN : ATL 72 LP TS: ; Ife d
SME WE NESS TAO (SS ZA rel
ee
AV a ie
‘
ie
* sid,
iv 1
bay fi
‘
.
|
y y
s i] > ¥ ~
a * :
J nq ;
, .
His i .
4 *
i) t .
; Ey ' i
i u
- ¥
} ¥ ‘ ue 7 y
¥ - ‘ ahi
1 - i\
SS
5 m
/ ‘
y
a oS bi ‘
1 , :
ist
x
f hile
aa x) Ls
DY
¥ rh
A =
in ¥ 4 . 7
‘ fi 4
i. 4 > 7 " ;
. Ven : ‘
Y a
i
Ki ba
: E i ; 4
* a tb 7
| os ft "
} ia) .t
i
Bert
i.) 5
AY ‘
' ‘7 ‘
yee | 4 "HH ‘
, & (AU UO
as it : :
cf i
1902. ] BUTTERFLIES OF THE FAMILY LYCHNIDA, 139
from some of the Pacific Islands, such as New Britain and the
Solomon group. The dominant section of the group is the genus
Arhopala, which alone contains over two hundred species and
varieties. Until the year 1890 these had generally been described
under the genus Amblypodia, in spite of the fact that they are
structurally quite distinct, and that Boisduval had created the
genus Arhopala, and had been followed in its use by the brothers
Felder in their important work in the ‘ Novara-Reise.’ Bois-
duval, however, omitted to diagnose his genus; and it has only
been since de Nicéville’s third volume of the ‘ Butterflies of India’
was published that the name has been generally used, but now
most students of the lLiycenidee adopt it. Mr. Moore has
attempted to split it up into six different genera, viz., Varathura,
Nilasera, Panchala, Satadra, Darasana, and Acesina; but none
of the characters he gives are constant, and though the markings
of Acesina are peculiar, yet the whole pattern follows precisely
that obtaming in Arhopala, and structurally they are identical ;
so that under these circumstances I see no object in retaining
that genus as distinct. The other genera also I am obliged to
disregard, none of them having so good a locus standi as Acesina.
The genus Arhopala forms one large homogeneous whole, and,
with the exception of one or two extreme forms, the merest tyro
would at once recognize them. I cannot therefore see what object
is gained by endeavouring to split them up into several genera
merely because of the large size of the genus. I retain Mr. Moore’s
genus Wahathala, which is very well marked ; but, with all respect
to that veteran observer, I cannot see my way to accepting the
other six.
I divide the whole group into six genera, viz., Amblypodia
Horsf., Zraota Moore, Surendra Moore, Thaduka Moore, Maha-
thala Moore, and Arhopala Boisduval. These Butterflies are very
sluggish in their habits, seldom flying unless disturbed, and then
only for a short distance, settling again as soon as possible on the
underside of the leaves, with which they assimilate well. They
prefer the thick forests, and but little is known of their life-
histories, though some of the eggs have been described by Doherty
and one or two other observers. Distant remarked on the shade
of blue varying considerably “inter se” at times ; and this I have
found to be the case, though I am unable to offer any real expla-
nation. I believe it will be found to be owing to some chemical
change, for it happens with apparently quite fresh specimens.
The colours of the whole group are aniline: in the majority of
the species they could not be made up, I believe, from the colours
of the spectrum, and it is therefore possible that they may be
more liable to change than otherwise.
In a dominant genus such as this is, we might look for certain
forms to be highly specialized in some way or other; and we find
that this is so, for we have a small number of species of that
brilliant metallic lustrous green colour that is so prominent a
feature in the Chinese and Japanese sections of the Thecline.
10*
140 DR. H. LYSTER JAMESON ON THE [ Mar. 4,
In one genus all the females are blue, thus proving that this
beautiful green development is probably of a comparatively
recent date. M. de Nicéville tells us that when seen in sunlight
these green specimens are gems of beauty, compared with which
the most brilliant of the blue species are absolutely dull.
Among many entomologists who have been most kind in
lending me their insects, I must mention two in particular who
have passed to the great majority. The late Dr. Staudinger sent
me over a considerable portion of his collection of this group,
including all his types; whilst the late M. de Nicéville (im whom
India has lost a most energetic and untiring observer) was good
enough to lend me the whole of this portion of his collection; so
that these two gentlemen alone furnished me with well over two
thousand specimens. My warm thanks are due not only to these
but to many others for much kind help.
This Memoir will be printed entire in the Society’s ‘Trans-
actions.’
The following papers were read :—
1. On the Origin of Pearls.
By H. Lyster Jamuson, M.A., Ph.D.
[Received February 7, 1902. ]
(Plates XIV. —XVII.* and Text-figures 22-24.)
Most theories of Pearl-production have assumed that the
‘“‘ nucleus,” whatever its origin may be, is the direct cause of
the secretion of a true pearl, and that the latter arises as a result
of the molluse’s endeavour to coat with carbonate of lime an
irritating body.
I do not propose in this paper to give yet another complete
historical survey of the various hypotheses, dating back to the
time of Pliny, which have been propounded. ‘These theories
have been summarized over and over again by writers on pearl-
formation. The more recent ones may conveniently be grouped
under the following heads:—That pearls are (1) concretions of
shell-forming fluid (Réaumur, 1717); (2) shell-substance deposited
around bodies or concretions of internal origin (Kiichenmeister,
in part, 1856; von Hessling, 1858; Pagenstecher, 1858) ;
(3) formed around an abortive or displaced ovum (Home, 1826 ;
Kelaart, in part, 1857); (4) secreted to coat over a grain of sand ;
(5) the result of injury to or perforation of the shell; (6) caused
by a parasite (Filippi and others); (7) formed in an ampulla in
the tissues (Hessling, 1858; Diguet, 1899). Several writers have
allowed the possibility of two or more of these causes.
The origin of the “ grain-of-sand ” theory is veiled in obscurity.
1 For explanation of the Plates, see p. 165.
1EX ZS) NOLS Waelbe luteal. 2CIGY.
EMUrda!. Bale & Danielsson E hith.
JAIMIE SON OI WBE OCRUGIUN Ol IRE ARULS .
DP ZS MOD wel el SCy,
6
ls
fa
~- 3a
Bale & Danielsson Lt dith.
ELL.J. del.
AWE SOI QIN AV Talla, NRG GIN, OIF, Te Neh AN ISUIE SS
Ey iS UGOZA oils del. SOWA,
H.L.J. del . Bale & Danielsson Lt¢ lith.
JAMESON ON THE ORIGIN OF PEARLS.
PZ. 8. WOZ, welll, IPL SCviil,,
v
Photo. mic.HAB. Bemrose & Sons, [!4 Collo.
JANIESON ON Waly OIRUGIONE OF Iie ANSUUisy
aa Y: n
fi. wal.
PRE) Gi vk '
‘
oy
ih iy Moh Gee.
1902.] ORIGIN OF PEARLS. 141
It has had many supporters, and still maintains a prominent
position in zoological text-books and popular compilations. It is
doubtless largely due to a confusion of true pearls with “blisters ”
or pearly excrescences on the shell. There is no recorded instance
of an undoubted sand-grain having been found in a pearl,
although hundreds have been examined. All attempts to produce
pearls by introducing such bodies into the tissues or between the
shell and mantle have led, at best, only to the formation of
“blisters.” Such methods of obtaining the latter have long been
known to the Chinese, and have repeatedly been applied in other
countries. Chemnitz, Beckmann, and others (1791) regarded
Linneus’s “ secret process” as merely boring the shells. However,
no subsequent boring experiments have yielded anything but
blisters, and the popular notion of Linnzus’s modus operandi is
little more than a guess. A great step in the right direction was
made when Filippi, in 1852, discovered the connection between
pearls and the presence of Distomum duplicatwm in Anodonta.
Filippi regarded these Trematodes as encysted. In his later
papers he allowed other forms such as Atax ypsilophorus to be
occasional causes of pearl-formation. He recognized that the
action of these parasites was specific, and compared it to the
formation of plant-galls. Kiichenmeister (1856) associated pearls
in Margaritana margaritifera with the larve of Atax ypsilo-
phorus van Beneden, which occur in the mantle, enclosed in cysts
secreted by the mollusc. He held that other parasites, as well
as bodies of internal origin, might also cause pearls.
Mobius (1857) found Trematode remains in pearls from the
Pearl-Oyster of the West Coast of America (probably Margaritifera
margaritifera L., var. mazatlanica Hanley’). Kelaart (1859)
held that parasites played an important part in pearl-formation
in Margaritifera culgaris (Schumacher) in Ceylon, but did not
associate any definite organism with it, although he found several
species living in the Pearl-Oyster. Thurston (1894) confirmed
the existence of platyhelminthan parasites in the same species,
but did not assert that they had anything to do with pearl-
production. Garner (1871) found that pearls in Mytilus edulis
and Margaritana margaritifera were due to Distomids, against
which the molluscs protected themselves by coating them with
calcium-carbonate. Comba (1898), who claims to have discovered
a method of producing free pearls by artificial means, says (p. 6)
that the cause is ‘un parassito il quale viene dal mollusco avvi-
luppato di strati di una bava che indurendosi forma la perla
formando cosi una pustola ed una pallina che cresce in grossezza.”
Dubois (1901) found in Mytilus edulis that the production of
pearls was due to Distomid larvee, to which (without description)
he applied the name Distomum margaritarum. His account of
the “ désagrégation” of formed pearls, and the liberation, to repeat
their life-cycle, of the parasites that form their nuclei, is quite at
1 Kor revised nomenclature of the Pearl-Oysters, see Jameson, 1901.
142 DR. H. LYSTER JAMESON ON THE [Mar. 4,
variance with my experience; and apparently presupposes that
the Trematode can survive complete calcification, which would
indeed be a very remarkable biological phenomenon. According
to Dubois it is only certain pearls that, by the death of the
Distomum, escape this annual disintegration and so reach greater
dimensions.
Von Hessling (1858), it seems, was the first to ascertain that
the pearl is formed inside an epithelial sac, and he emphasized
the importance of this structure. He regarded the sac as being
derived from the blood-cells.
This sac has been noted by Diguet (1899), who suggests that
it may be due to the stimulation of a parasite. I can find no
support for Diguet’s view, that the formation of the pearl in this
sac proceeds on different lines to those on which the substance of
the shell is deposited.
The “vesicle or bag of the ovum” figured by Home (1826,
pl. xiii.) may also be this sac.
Before entering upon an account of my own observations, I
wish to express my thanks to Mr. H. H. Arnold Bemrose for
kindly preparing the microphotographs which accompany this
paper; to Baron Louis d’Hamonville for much valuable inform-
ation concerning the pearl-bearing mussels of Billiers; to
My. A. Scott, of the Lancashire Sea-Fish Hatcheries, for supplying
me with abundant material from the Piel mussel-beds; and to
Mr. W. Wells, Marine Superintendent at the Brighton Aquarium,
for conducting experiments for me.
The distribution of pearl-producing individuals of Margaritifera
margaritifera L., M. maxima Jameson, Pinna mgrina Lam.,
ippopus hippopus L., and Tridacna gigas Lam., in New Guinea
and Torres Straits, suggested to me that pearls were the result
of a specific pathological condition, and that the circumstances
necessary to ensure infection were present only in certain areas,
often of small extent. I soon convinced myself, by a study of
material that I brought home with me, that Trematodes formed
the nuclei of some of the pearls in each of the above-named
species, but that others contained nothing more than a few
yellowish granules in the centre. The same results were obtained
with specimens of M/ytilus edulis, sent me from Lancashire by my
friend Mr. James Johnstone. In all cases where the pearls had
been preserved in stéw in the tissues, they were found to be
enclosed in a sac composed of an epithelium physiologically and
histologically identical with the outer shell-secreting epidermis of
the mantle. This observation at once accounted for the similarity
in structure between the layers of the shell and those of which a
pearl iscomposed. The obvious conclusion was that this sac is the
direct, and the Trematode the indirect, cause of pearl-production,
and that the key to the problem of the origin of pearls might be
obtained by investigating the origin of the sac and its relations
to the Trematode.
1902.] ORIGIN OF PEARLS. - 143
As it was not possible for me to return to the habitat of the
true Pearl-Oysters, I selected the Common Mussel (Mytilus edulis)
as a suitable species upon which to begin my observations. This
molluse produces pearls in many localities on the coasts of Europe,
but it is only on certain beds that pearls are abundantly formed.
The most favourable places seem to be estuaries or land-locked
channels. In such situations pearls may be found in almost
every example, except those which are attached to stakes or
floating objects, and so raised off the bottom.
The pearls produced by the Common Mussel are, like the
nacreous lining of the shell, lacking in lustre. They are generally
white or silvery, but blue and brown examples are not uncommon.
They have no value as gems, though, strange to say, a market
seems to have existed for them in the first half of the last century
(“ D.C.,” 1830).
They are mostly formed in the subcutaneous tissue of the
dorsal body-wall or in the mantle-lobes. When they occupy a
more deep-seated position they have probably been secondarily
displaced.
In the little harbour of Billiers (Morbihan), situated on the
estuary of the Villaine, there is a colony of pearl-bearing mussels,
that has been described by d’Hamonville (1894). After reading
d’Hamonville’s account, I was struck with the idea that this
colony should offer special facilities for investigating the causes of
pearl-formation and the conditions for infection. D’Hamonville
found that although the mussel is abundant all round the coast,
pearls are only produced in the harbour itself, the beds being, at
most, only a few acres in extent. Here almost every shell, if not
too young, contains pearls.
I visited Billiers in August 1901 and again in December of the
same year, and had no difficulty in findmg the parasites and
tracing the part played by them in pearl-formation. They were
the larve of a Distomid belonging to the subgenus Lewcitho-
dendriwm (Loos), and very closely resembling Z. somaterice
(Levinsen), which in the mature condition inhabits the intestine
of the Eider Duck. I found larvee, very similar to these, in Sporo-
cysts in Tapes decussatus, and subsequently proved the infection
of Mytilus experimentally from these Sporocysts. In September
of the same year I visited Piel, Lancashire, and found that there
also pearls are caused by the same parasite, but that in this case
Cardium edule acts as “ first host” for the Sporocyst.
Finally, in December 1901, when I revisited Billiers, I examined
five specimens of the Common Scoter or Black Duck, Wdemia
nigra L., which is notorious in the Villaine for its depredations
on the mussel-beds, and is locally called, on account of its habit
of feeding on Mytilus, “‘Cane mouliére.” Every one of these
specimens was teeming with Distomum (Leucithodendrium)
somaterice.
For histological work, pearls were decalcified im situ in the
tissues and then sectioned. Others were decalcified, cleaned, and
144 DR. H. LYSTER JAMESON ON THE | Mar. 4,
examined whole; while others again were ground down on a
Water-of-Ayr hone, care being taken that the “ Schliff” so made
should pass as nearly as possible through the centre. The
Cercaric and Sporocysts were either examined entire, or sectioned
in situ.
For decalcifying, it was found best to use pearls preserved in
spirit, as those that had been kept in a dry state, although often
giving good results when examined entire in oil of cloves, were
unfit for cutting with the microtome, owing to the brittle and
horny nature of the dry conchyolin. The most satisfactory effects
were obtained by using very dilute (‘5 to 1 per cent.) nitric acid
in water. Stronger solutions often caused bubbles of carbon
dioxide to be evolved in the residual conchyolin, but by using
these weak solutions freely the gas was dissolved almost as soon
as formed. The time required for this process varied from a few
days to some weeks, according to the dimensions of the,pearl and
the proportion of conchyolin to salts.
Structure of the Mantle and Shell.
The tissues composing the mantle of Mytilus edulis are :—
(1) An external simple epithelium, which is a direct continua-
tion of the dorsal body-wall ;
(2) An internal ciliated epithelium, resembling the epidermis
of the foot and gills; and
(3) A spongy connective tissue; in the meshes of which the
blood circulates.
The external epithelium (Pl. XIV. figs. 1-4, Pl. XV. fig. 5,
eat.ep., and text-fig. 22) is composed of a single layer of flattened
columnar cells with spherical or ovate nuclei. The outer surfaces
of these cells are closely applied to the inner nacreous substance of
the shell. The appearance of this epidermis differs considerably
according to the degree of contraction and method of preservation.
Its constituent cells are polygonal in surface view, and brick-shaped
or columnar in sections at right angles to the surface.
Their protoplasm stains rather more strongly with hematoxylin
than that of the connective tissue, and shows faint strie perpen-
dicular to the surface. Their bases are attached to the connective-
tissue fibres. Scattered here and there throughout this epithelium
are spherical cells which stain lightly. They may be the “ Hirund
kérnige Zellen” that Tullberg describes (1882). They are never
numerous, and I am inclined to attribute their presence largely to
defects in preservation.
The epidermal lining of the mantle-cavity (Pl. XIV. fig. 2,
Pl. XV. fig. 5, and text-fig. 22, int.ep.) is the typical glandular
ciliated epithelium so common in the skin of invertebrates. It
is composed of columnar ciliated cells and interstitial gland-cells,
some of which project basally into the connective tissue. It is
1902. | ORIGIN OF PEARLS. 145
altogether a much more heterogeneous layer than that applied to
the inner surface of the shell, and is usually rather thicker.
The connective tissue (Pl. XIV. figs. 1-4; Pl. XV. fig. 5, ¢.t.)
is a meshwork of irregular or stellate cells with oval or spindle-
shaped nuclei, and more delicate fibres the nuclei of which are
relatively longer and narrower, and stain more deeply than those
of the former. There is a perfect intergradation between the two
kinds of cells, and their relative abundance varies in different
individuals according to the condition of the gonads. The fibres
are more numerous just under the epidermis than elsewhere. The
blood circulates in the spaces between these cells and fibres, and
in places these lacunz are enlarged to form regular blood-sinuses.
Numerous blood-corpuscles (/.) can be seen in the meshes of the
connective tissue, especially under the outer and inner epidermal
layers.
Yellow refractive granular masses, showing traces of cellular
structure, sometimes occur in the meshes of the mantle paren-
chyma, especially in old mussels. These may be the remains of
the broken-down gonads of previous years, or groups of dead
leucocytes.
In the connective tissue are embedded the nerves and muscle-
fibres of the mantle, and the gonads when ripe extend into it.
The structure of the shell has been very thoroughly investi-
gated by von Nathusius Kénigsborn (1877), Tullberg (1882), and
Khrenbaum (1885), to whose observations | can add nothing new.
The method in which the shell is laid down is of great interest
on account of the identity in structure between the substance of
pearls and that of the shell. Biedermann’s recent paper (1901),
which is full of new and significant facts and carefully summarizes
previous observations, proves beyond all question that the organic
basis of the shell (conchyolin), which is present also as the basis
of pearls, is a true cuticular product, secreted or excreted by the
underlying epidermis of the mantle. Biedermann shows that in
both Lamellibranchs and Gastropods the calcareous substance of
the shell can only be deposited in such a cuticle.
The cuticular conception of the conchyolin was, I believe, first
propounded in this form by Huxley (1859). In sections of the
decalcified shell and mantle, I find that the epithelium is generally
applied closely to the conchyolin, and its cuticular outer surface
is apparently directly continuous with the latter. If, during the
process of fixing, the mantle has been separated from the shell, a
certain amount of uncalcified conchyolin may be found attached
to the epidermis. Moreover, if the mantle of a live mussel be
carefully stripped from the inner surface of the shell, a delicate
transparent membrane, like that which Huxley found in Anodonta,
but less conspicuous, can be detected. This membrane tears away
irregularly, some parts adhering to the mantle, others to the shell.
This irregular tearing is a further evidence that the uncalcified
membrane is in continuity with both shell and mantle. That the
mantle can, however, detach itself from the inner surface of the
146 DR. H. LYSTER JAMESON ON THE [ Mar. 4,
shell is obvious in such forms as Margaritifera, Pinna, and
Tridacna, where the mantle-margin is freely retractile.
The evidence adduced in support of the alternate theory of
“growth by intussusception, originated by Mery (1712), revived by
von Nathusius (1877 & 1898), and supported by Felix Miiller
(1885), is not convincing. The facts supposed to lend weight to
this hypothesis are quite explicable on the apposition theory.
Pearls, Blisters, and Coneretions.
As some confusion exists as to the exact connotation of the
word pearl, I propose to adopt in this paper the terms “ pearl,”
“blister,” and “free concretion” for three different kinds of
structures that occur in molluscs.
Pearls.—A pearl consists of one or more layers of shell-substance
(é. e., conchyolin in which the crystals of inorganic matter are
disposed in the same manner as in the shell), enclosing a central
nucleus, and formed in a closed sac embedded in the tissues.
This sac is composed of epithelial cells similar to those that form
the outer mantle-epidermis. This sac is first formed around a
parasite, which probably exercises a specific stimulation.
In Mytilus edulis and many other forms this parasite is a larval
Trematode, but it is probable that certain other parasites can
stimulate some molluscs to form such sacs. The parasite does not
necessarily become the nucleus of the pearl, but may escape from
the sac before calcification.
Any of the substances which form the different parts of the shell
may be represented in a pearl. ‘Thus we have nacreous pearls,
prismatic pearls, the periostracum pearls of Modiola modiolus
formed in the mantle-margin, pearls a part of which may be
formed of the transparent striated substance which characterizes
the attachment of the muscles to the shell, and pearls formed
entirely or in part of the substance of the hinge-ligament. Large
brown leathery hinge-pearls are occasionally found in Torres
Straits in Margaritifera maxima Jameson.
A pearl may become secondarily fused to or embedded in the
substance of the shell by the absorption of intervening tissues
(text-fig. 22). These pearls are sometimes spoken of as attached
pearls. Similarly two or more pearls may become fused together,
forming double or compound pearls, of which a notable example
is the celebrated “ Southern Cross.”
The various substances, when two or more are present, are
not always arranged exactly in the reverse order of the layers
of the shell, as sometimes stated. We may, indeed, have several
alternations of nacre and prismatic substance, or of the latter
and conchyolin. The kinds of shell-substance entermg into
the composition of a pearl are determined by the position of the
latter. De Villepoix (1892) has shown that different parts of the
mantle-epithelium are concerned severally in the formation of
periostracum, prismatic and nacreous substance. Obviously the
1902. ] ORIGIN OF PEARLS. 147
epithelium of the pearl-sac acquires the special characters of the
adjacent part of the epidermis.
Blisters.—It is proposed to confine this term to internal
excrescences of the shell, which are caused by the intrusion of
foreign bodies between the mantle and the shell, or by the secre-
tion of a nacreous cicatrix to close the perforations of boring
molluscs, worms, or sponges. These are sometimes referred to
as “attached pearls” or even as “pearls,” but have a totally
different mode of origin and should never be confused with the
latter.
Coneretions.—In many molluses small free calcespheritic bodies
occur at times in the connective tissues, which, not being enclosed
in epidermal sacs, cannot acquire the structure of the shell-
substance. They are probably due to different causes in different
molluscs. In Yapes they are frequent, and are due to the
calcification of degenerated Sporocysts or of dead Cercariz con-
tained in the same. Similar concretions, which I found in
Pholas candida at Billiers, were caused by dead Cercariz of
another species, contained in Sporocysts.
Old examples of Mytilus edulis L., Modiola modiolus I..,
Hippopus hippopus l., Margaritifera vulgaris (Schumacher), and
Anodonta sometimes contain similar bodies, but their origin in
these cases is uncertain.
In all instances that have come under my notice they are more
or less spherical, and composed of needle-like prisms of carbonate
of lime radiating from a centre.
Structure of Pearls.
A. Mytilus-pearl examined entire often shows a darker spot in
the centre, which corresponds to the ‘‘nucleus.” The nucleus is
always visible in a section ground from the pearl, though its size
varies from ‘1 mm. to ‘7 mm.
It is often yellowish brown or black, the colour being imparted
by the dead remains of the Trematode, or by the small amount of
residual matter left, if the worm has escaped from the sac
(Pl. XVII. figs. 12-16). The crystalline structure of the nucleus
is quite different to that of the remainder of the pearl and to that
of the shell. We find in the nucleus one (Pl. XVII. fig. 16) or
more (Pl. XVI. fig. 8, Pl. XVII. fig. 14) centres of calcification,
consisting of spherical masses of radially arranged crystals. Each
centre of calcification, if more than one be present in the nucleus,
shows a distinct black cross when viewed between crossed nicols.
Sometimes the nucleus is irregularly or incompletely calcified
(Pl. XVII. figs. 14-16). The resemblance which concretions
and the nuclei of pearls bear to Harting’s bodies (Harting, 1872)
is interesting; for they are formed, so far as we can judge,
in a similar manner, namely, by the slow precipitation of
carbonate of lime in a viscous substance like albumen or decaying
animal matter. On the other hand, the peripheral parts of the
148 DR. H. LYSTER JAMESON ON THE [Mar. 4,
pearl are, like the shell, formed by the calcification of the cuticle
of the living cells, and owe their structure to the special characters
of that membrane or of the underlying epidermis.
A section of a decalcified pearl shows the nucleus, in which the
cuticle and sometimes the suckers of the Distomum can be
distinguished. Occasionally the outlines of the soft parts (e. g.,
pharynx and digestive ceca) are still visible, as in Pl. XIV. fig. 1,
ph. & dig. More generally, however, nothing can be seen but a
mass of yellowish-brown granular substance surrounded by the
cuticle (text-fig. 22).
There is often a certain amount of refractive granular matter
associated with the remains of the worm, probably an excretion ;
and, if the parasite migrates out of the sac, this may form the
inconspicuous nucleus of a pearl.
Just as the peripheral parts of a pearl present, when ground
down to a thin section, a similar structure to that of the shell,
so the conchyolin basis of a decalcified pearl shows the same
characters. The outermost layer of the latter is uncalcified and
continuous with the cuticle of the cells of the sac, just as the
outer mantle-epidermis is attached to the inner surface of the
shell (Pl. XIV. fig. 1, con.').
There is no organic union between the conchyolin and the
nucleus.
The sac containing the pearl is composed of a simple columnar
epithelium (Pl. XIV. fig. 1 & text-fig. 22, s.), which in its histo-
logical structure, as well as in its power of secreting as a cuticle
the conchyolin basis of the pearl, is indistinguishable from the
outer epidermis of the mantle.
Blood-spaces, containing corpuscles, are well developed around
the sac.
Such a pearl cannot then be compared—as some writers have
suggested—with the concretions or calculi of cholesterin or other
substances found in the vertebrate body, but rather with the
structures sometimes found in epidermoid tumours and atheroma
cysts.
Origin and Development of the Pearl.
The Trematode enters Mytilus edulis as a tailless Cerearia, and
at first may often be found between the mantle and shell. It is
probable that it reaches this position by boring through the
mantle, but I have not yet been able to find one in the act of
doing so. The larvee creep about on the inner surface of the
shell, and, after a while, again enter the connective tissue of
the mantle, where they come to rest, assuming a spherical form.
They seem to avoid the more muscular parts of the mantle—no
doubt because the absence of a definite boring apparatus makes
it difficult for them to pass through the latter. When embedded
in the tissues they are visible to the naked eye as little yellowish
spots, about 3 mm. in diameter.
At first the worm only occupies a space lined by connective-
1902. ] ORIGIN OF PEARLS. 149
tissue fibrils (Pl. XIV. fig. 2), but soon the tissues of the host give
rise to an epithelial layer, which lines the space and ultimately
becomes the pearl-sac (Pl. XV. fig. 5, s.).
This epithelium appears to arise quite independently of the
outer epidermis, and is no doubt due to a specific stimulation on
the part of the parasite, as other parasites, e. g. Sporocysts, Cestode
larvee, &e., are not surrounded by such a sac.
At first a few cells appear (Pl. XIV. figs. 2, 3, pr.), which pro-
liferate and arrange themselves along the walls of the cavity.
These cells are larger than the connective-tissue corpuscles, and
more susceptible to stains. They are flattened and polygonal in
surface view. Their nuclei (Pl. XIV. fig. 3, 7.) are large and
spherical, and show the conspicuous chromatin reticulum and
distinct nucleolus that characterize the nuclei of embryonic or
rapidly dividing tissues.
I have not been able to find the nuclei of these cells actually
undergoing division. The proliferating sheet of cells ultimately
surrounds the parasite and becomes the sac. From the first
these cells are basally continuous with fibres of connective tissue
(Pl. XIV. fig. 3, ¢.¢.). Their transformation into the pearl-sac is
a gradual one, and every step can be traced in sections of the
parasites @ site.
If the Trematode larva completes its maximum possible term
of life it dies, and the tissues of the body break down to form a
structureless mass, which retains the form of the parasite owing
to the rigid cuticle. :
Tn this mass arise one or more centres of calcification (Pl. XVI.
fig. 8), and the precipitation of carbonate of lime goes on until
the whole larva is converted into a nodule which has the calco-
spheeritic structure already described for the nucleus. The
granular matter surrounding the worm, if present, also undergoes
calcification.
The epithelium of the sac then begins to shed a cuticle of
eonchyolin (Pl. XIV. fig. 1), and from this point the growth of
the pearl probably takes place on the same lines and at the same
rate as the thickening of the shell.
The sac sometimes begins to form pearly substance before the
worm is completely calcified (Pl. XVII. fig. 16).
The Distomid larve sometimes leave the sac formed around
them, and voluntarily migrate into other parts of the body
before again settling down. Empty sacs may be found in the
mantle, and old specimens of the larva (distinguishable from
recently immigrated ones by their darker colour and laden
excretory organs) sometimes occur free between the mantle and
the shell.
The occurrence of pearls in which the nucleus is not a Trematode
but merely a few refractive granules (Pl. XVII. fig. 13) can be
accounted for in this manner.
Some compound pearls are evidently formed by short migrations
on the part of the Cercariz, which leave a small amount of
150 DR, H. LYSTER JAMESON ON THE [Mar. 4,
residual material in the sac, vacate it, and settle down in the
immediate vicinity (Pl. XVII. fig. 15).
The residual matter in the first sac forms the nucleus of a pearl,
and if the Trematode dies another is formed in the new one beside
it. If these pearls grow and fuse a double pearl is formed, the
nucleus of one half being obviously a Trematode, that of the other
being merely granular matter.
Thave traced three stages in the formation of such a pear]—the
first, in which a Cercaria is found in a sac with a vacated sac close
by ; the second, in which a small pearl is close to the live Cercaria ;
and the third, in which two or more small pearls lie close together,
only one having a Trematode for nucleus.
Dubois (1901) suggests that the death of the Distoma may
sometimes be determined by Sporozoa, some members of which
group are known to attack Trematodes. In one of the specimens
that I sectioned there was a parasitic protozoon embedded in the
tissues. If such parasites were to occur frequently they would
of course facilitate and intensify the production of pearls. But
they are not essential, any more than the presence of the dead
Distoma in the sac is necessary for pearl-formation.
Text-fig. 22.
eExXtiep. Com ns. Ss
co,
Le
Sue
Le
bu
OAS : Wy, &
Po YS y
US Nee
Z| Pe pie mee a it
OS eka 1@5a% A
OS,
D6 DOO! OF PEC JO JORGE 25 |0/ eS] elo
mM
int. ep.
A Pearl about to become attached to the Shell.
nu., nucleus of pearl ; int.ep., internal ciliated epidermis of mantle; eat.ep., external
epidermis of mantle; con., conchyolin basis of pearl; s., epithelium of pearl-sac ;
c.t., connective tissue. XX 50.
A pearl may increase in size until its diameter is considerably
greater than the thickness of the mantle, so that it protrudes
visibly. It may even break through the ciliated epidermis; for
valuable pearls have been found in the branchial chamber and
also outside the shells.
1902. ] ORIGIN OF PEARLS. 151
If it presses upon the tissues intervening between itself and the
shell, these may become absorbed, in which case the epithelium
of the pearl-sac becomes continuous with the shell-forming epi-
dermis (text-fig. 22). The result is that the subsequently formed
layers of the pearl are continuous with those of the shell, and an
attached pearl is formed. The fusion of two or more pearls to
form a compound pearl is effected in the same way.
Structure of the Trematode Larva.
With the exception of the female reproductive organs, which
are as yet undeveloped, the larva presents all the characters of
Distomum (Brachyceliwm Dujardin, 1845, Leucithodendriwm Loos,
_ 1896) somaterice (Levinsen, 1882), from the Eider Duck, Greenland.
(For sub-classification of Dujardin’s subgenus Brachyceliwm, see
Stossich, 1899.) The body (Pl. XV. fig. 6) is oval, blunter in
front than behind, and tapering markedly in the last third. In the
resting condition inside the sac it is nearly spherical. The average
dimensions are 55 mm. to "7 mm. The extreme sizes seen were
‘45 mm. and ‘75 mm. The oral sucker is larger than the ventral
one, the ratio of their diameters being usually about 4:3. But
in this point there is a considerable amount of variation both in
the larva and in the adult Z. somaterie. Except on the surfaces
of the suckers, which are smooth, the cuticle is beset with small
spines (text-fig. 23). These are arranged in transverse rows, the
Text-fig. 23.
Cuticle of the Cercaria, in surface view. X 700.
members of which also form diagonal rows, so that the cuticle
in surface view appears to be divided up into little diamond-
shaped fields. There are about two hundred transverse rows of
spines on the dorsum. Immediately around the ventral sucker
the spines occur in concentric circles. The connective tissue is
the typical parenchyma of the Flatworms (Pl. XIV. fig. 2; Pl. XV.
figs. 5, 7, pa.). It is richer in nuclei immediately under the skin
than elsewhere, especially on the dorsal surface (Pl. XV. fig. 7,
pa.n.). Some muscle-fibres are present in the connective tissue
running from the body-wall to the suckers and pharynx (Pl. XV,
fig. 7, m.f.). Theemusculature of the body-wall consists of an
outer circular layer (Pl. XV. figs. 5, 7, c.m.) immediately under
152 DR. H. LYSTER JAMESON ON THE [ Mar. 4,
the cuticle, a deeper longitudinal layer (/.m.), and, on the ventral
surface, a less defined tract of transverse fibres inside the longi-
tudinal muscular coat.
The suckers are lodged in a slight involution of the cuticle
(Pl. XV. figs. 5, 7). Their relative and absolute sizes in surface
view are determined by the degree of contraction of their con-
stituent fibres. The ventral sucker is about one-fourth or one-
third of the total breadth of the body.
The mouth is situated in the middle of the anterior sucker,
and generally appears triangular in sections (Pl. XV. figs. 6, 7, m.).
The funnel-shaped buccal tube opens behind, by a narrow orifice,
into the spherical muscular pharynx (ph.). This is followed by
the short straight cesophagus (@.), which, passing upwards and
backwards, bifurcates to form the sac-like digestive ceca (Pl. XIV.
fig. 2; Pl. XV. figs. 5-7, dig.), which are dorsal to the other organs.
In the resting worm these ceca are greatly distended with
yellowish granular material, doubtless derived from the tissues of
Mytilus.
Even the csophagus is often tightly crammed with food.
The digestive system in this condition occupies the bulk of the
body, anterior to the ventral sucker, but when empty is much
smaller. The posterior end of the pharynx is provided with a
group of salivary glands (Pl. XV. figs. 6,7, s.gl.). The epithelium
of the digestive system consists of very large flat polygonal cells
with conspicuous nuclei (Pl. XV. fig. 7, int.ep.). The individual
cells can sometimes be distinguished in pressure preparations.
There is an ill-defined supra-pharyngeal nerve commissure (Pl. XV.
fig. 7, n.c.) and a pair of lateral cords. The excretory system ~
(Pl. XIV. fig. 2; Pl. XV. figs. 5—7, ex.) consists of two enormous
tubular sacs, extending to the anterior end of the body and
converging to form a pyriform median vesicle, which opens by
a pore at the hinder end (Pl. XV. fig. 6, ex.p.). The excretory
tubes are generally quite full of opaque spherical granules,
presumably of excretory matter. When treated with hydro-
chloric acid they become transparent (Pl. XV. fig. 6).
In living specimens a few flame-cells can be seen in short,
lateral, and apparently unbranched tubules given off by the
excretory sacs. But the distended condition of the latter makes
it difficult to ascertain their precise relations.
The female organs are not developed in the resting larva.
The worm is protandrous, and the male genital organs reach a
conspicuous size, even in the Sporocyst. The rudiments of the
testes, vasa deferentia, and penis are very obvious in sections
(Pl. XV. figs. 5, 7) and in stained preparations of the entire
worm (Pl. XV. fig. 6). Being composed of young cells they stain
deeply. In fresh specimens they are less obvious. The penis
opens out at the genital. pore (Pl. XV. fig. 7, g.p.), which is
situated immediately in front of the anterior border of the
ventral sucker. It is an elongated hollow pyriform body, lying
in front of and dorsal to the sucker. The rudiment of the
1902. ] ORIGIN OF PEARLS. 153
seminal vesicle receives the vasa deferentia (Pl. XV. fig. 6, v.d.),
which can be traced back into the spherical testes (¢e.).
Biology.
When the larva first enters Mytilus it is somewhat smaller
than the resting specimens and more transparent. The excretory
organs, which are laden with granules while in the Sporocyst, are
comparatively empty, and the gut is not yet distended with food.
As it grows older both the excretory and digestive systems become
more and more laden, so that they form the great mass of the
body (Pl. XIV. fig. 2; Pl. XV. fig. 5). It is largely to the
contents of the latter that the parasite owes its dark yellow
colour, the cuticle being pale golden or straw-coloured.
The worm often excretes some granular substance, which may
almost surround it in the sac. It is this stuff that serves as
“nucleus” for a pearl, if the Trematode migrates to another part
of its host.
It is interesting to note that at no period is this worm encysted,
in the sense in which the Liver-fluke and so many other
Cercarie encyst. The dark colour of the epithelial sac, which
can often be isolated with the worm, suggests, on casual obser-
vation, that the latter is encysted, but I have determined by
sections that this is never the case.
In a certain sense it is a resting stage, but the distension of
the alimentary system makes it obvious that it is also a highly
assimilative phase in the life of the worm, which is storing up
energy for the maturation of the gonads, on reaching the final
host.
Life-history of the Parasite. The Sporocyst Stage.
After a laborious examination of most of the organisms which
inhabit Billiers Harbour, I was rewarded by finding, in Tapes
decussatus Gmel., Sporocysts containing tailless larve, almost
identical with those which occur in Vytilus (Pl. XVI. figs. 9, 10,
and text-fig. 24). They differed from the latter only in their
smaller size, paler colour, more distended excretory organs and
empty gut, and in the possession of special sense-organs and eyes.
Tapes decussatus is extremely abundant in Billiers Harbour,
living in burrows about six inches deep in the black gravelly
clay that forms the bottom. The local name of this mollusc is
Palourde, and it is extensively collected for food. I am informed
that although Tapes occurs in small numbers all along the shores
of the Villaine, it is nowhere found in sufficient quantities to be
worth fishing, except in Billiers Harbour. Indeed, when I visited
Billiers in December 1901, a lugger from the other side of the
estuary came over specially to collect this shell and Mya arenaria
for the markets, there being no supply on the south shore.
I found Sporocysts in every specimen of Tapes that I examined,
numbering nearly two hundred. They occur in the muscular or
Proc. Zoou. Soc.—1902, Vou. I. No. XI. 11
154 DR. H. LYSTER JAMESON ON THE [ Mar. 4,
connective tissue of the mantle-margin, where it is attached to
the “pallial line,” especially along the ventral border, and
around the insertion of the siphonal musculature. The favourite
place seems to be the dorsal side of the latter. The individual
Sporocysts are embedded in and closely adherent to the bundles
of muscle-fibres.
In young examples I found small, simple, spherical or oval
Sporocysts, about *5 mm. in diameter, and containing 6-10
Cercariz, but in larger examples groups of Sporocysts occur
(Pl. XVI. fig. 9, and text-fig. 24).
Text-fig. 24.
Tapes decussatus.
Group of Secondary Sporocysts, as seen in a pressure preparation.
m.é., muscular tissue of Tapes; sp.c., Sporocyst; cer., Cercaria contained
in Sporocyst.
The individual Sporocysts in these cases are often larger than
the simple ones. The groups are, from their position, their
relations to one another, and their progressive increase in size
and number of constituent cysts as Z'apes grows older, evidently
produced by budding or secondary division of the original simple
ones. Their growth is very slow, and their duration of life must
be practically co-extensive with that of their host. In large
specimens of Zapes they appear to represent several successive
infections; as we may have, in the same individual, large groups,
probably several years old, smaller ones containing three or four
cysts, and finally little single Sporocysts like those found in young
individuals. On the other hand, in young Zapes, 10-20 mm.
long, although the simple cysts and small groups occur, the large
groups are not to be found.
Specimens of Zapes measuring 12x10 mm., with one marked
period or interruption in the growth-rings on the shell, contained
1902.] ORIGIN OF PEARLS. 155
small simple Sporocysts, or, at most, groups of three or four
secondary ones. Examples which averaged 17—21 mm. in length
contained masses of seven to ten cysts, those measuring 27 mm.
had still larger groups, while from that size upwards there was a
steady progression in the dimensions and number of constituent
units in the groups of Sporocysts. Some of those found in Z'apes
40-50 mm. long measured 7 mm. in diameter, and contained as
many as thirty secondary cysts, and a hundred Cercariz or even
more.
As the Sporocyst grows, it sometimes compresses the tissues
that intervene between it and the shell, which apparently inter-
feres with the secretion of fresh shell-layers. This leads to the
development of white chalky spots on the inner surface of the
valve. These patches, in old individuals, further prove that the
large groups of Sporocysts are the descendants of the original
small ones which are present when the molluscs are young. For
we may see two or more white scars on the lining of the shell,
marking the position of the cysts when the shell was younger.
In fact, the Sporocysts may leave, imprinted on the shell, the
history of their movements as the mantle-margin shifts outwards,
just as the adductor muscles mark on the nacre the record of their
migrations. The first or innermost of these scars is a small one,
such as might result from a triple or quadruple cyst, the next
is larger, while the group of Sporocysts in the mantle-margin is
larger again. This plainly shows that as the mantle-margin
followed the peripheral growth of the shell, the group of Sporo-
cysts increased in size.
These compound Sporocysts are, of course, thicker than the
normal thickness of the mantle, and stand out as opaque white
granular eminences, obvious as soon as the shell is opened.
On the beds of Pearl-bearing Mussels in the Barrow Channel,
opposite the Piel Fish-Hatchery, where every specimen of Mytilus
is abundantly infected with the Leucithodendriwm, and almost
every specimen contains pearls, Tapes is not found. The Cockle,
Cardium edule L., is common there, and acts as a host for the
Sporocysts. Somewhat less than half the specimens of Cardiwm
that I examined at Piel were infected. In Cardiwm the Sporo-
cysts occurred in the mantle-margin, close to the anterior border of
the anterior adductor muscle. Large groups, such as occur in
Tapes, were not observed, but only single, triple, or quadruple
cysts.
I have not yet been able to trace the infection of Tapes or
Cardium. It is therefore impossible to say whether infection
takes place by means of a free swimming Miracidiwm larva or not.
The constant occurrence of the Sporocysts in exactly the same
positions suggests that the eggs are carried into the digestive
system of Zapes with the food-bearing current, hatch out in the
alimentary canal, enter the circulatory system, and reach their
destination via the posterior pallial artery, along the course of
which they are distributed in Tapes. Moreover, ae ee in
156 DR. H., LYSTER JAMESON ON THE [ Mar. 4,
which I found the Sporocysts in Cardiuwm at Piel corresponds to
the end of the anterior pallial artery.
The Cercaria in the Sporocyst first appears as a little oval
cellular ball, budded off from the wall, measuring about ‘05 mm.
During the early stages of its development it is transparent, and
its structure can be made out without difficulty ; but, as it grows,
its excretory organs become gradually laden and distended with
opaque granules, which conceal the other parts.
The fully formed Cercariz in the Sporocysts measure "15 mm.
to'3 mm. They are whiter than those found in Mytilus, but the
arrangement of the spines on the cuticle is the same. They
possess a pair of pyramidal or conical light brown eye-spots
(Pl. XVI. fig. 10, ¢.), each provided with a lens. There are about
six tactile papille (¢.p.) at the anterior end of the body. These
sense-organs are no doubt serviceable to the larva, during its free
living stage, after leaving Tapes, and before entering Mytilus.
The digestive system (Pl. XVI. figs. 9, 10, dig.) is empty, and
occupies less space than in the Mytilus stage. The form and
relations of the suckers and pharynx are the same. The penis
(Pl. XVI. fig. 9, pe.) and testes are already developed, and have
the same relations as in the Mytilus worm. In pressure prepara-
tions of the live worm they are not easy to discern.
The majority of Cercariz in the Sporocysts are fully developed,
young transparent ones being less common. They probably
remain a considerable time before vacating it. A few on their
way out may often be found free in the tissues of the mantle.
Search in the mud and with the tow-net at Billiers, failed to
reveal the free living stage. I have, however, found examples in
water in which Z’apes had been kept for some days.
This Trematode is not provided with a cercarian tail at any
stage of its existence, and it is only capable of creeping movements.
The larva in the Sporocyst is rather more active than the later
stage which occurs in Mytilus.
Tf a Cercaria dies while still in the Sporocyst, its remains
become calcified; but, not being enclosed in an epidermal sac,
secreted by the mollusc, it does not give rise to a pearl, but merely
to a concretion. Again, an exhausted Sporocyst may undergo
similar calcareous degeneration with the same result.
Artificial Infection of Mytilus.
On leaving Billiers in the beginning of September 1901, I
brought with me about fifty infected examples of Zapes. I first
placed these in a tank at the Piel Fish-Hatchery, which Professor
Herdman and Mr. Scott kindly placed at my disposal. In order
to test experimentally the infection of Mytilus from Tapes, I put
in the same tank about seventy mussels, taken from the piles of
the old pier at Piel. These mussels of which I examined a
number, were practically without parasites. About one in every
five of the largest examples contained a Cercaria, one had two
1902. ] ORIGIN OF PEARLS. 157
Cercarie, and one contained a small pearl. It is apparently
difficult for infection to take place, except on the bottom, owing
to the absence of swimming-organs in the parasite. Hence the
absence of Cercariz in these examples.
Eleven days after they were placed in the tank I examined two
of these Mytili, and found that the first contained one and the
second two Cercariz. These Cercarize were recently immigrated
examples, as they were small, rather transparent, and not yet
surrounded by sacs.
T then transferred the experiment to Brighton, where Mr. W.
Wells, the Marine Superintendent at the Aquarium, kindly kept
the molluscs in a tank in his private office.
On the 18th of November, 1901, two months after the specimens
were placed in this tank, I examined six of the mussels. Of these
one contained six Cercarize, another four, two had each three
parasites, one contained two, and one was still uninfected.
When the experiment was transferred to Brighton I added
about two dozen mussels that had been in the Brighton Aquarium
for two years. J examined six such mussels before introducing
the others, and found that none of them contained live Cercariz,
though four of them had one small pearl apiece.
On 5th April, 1902, I took up a sample of 10 mussels from this
tank, comprising five of the specimens originally taken from the
pier at Piel, and five of those that had been transferred from
another tank at Brighton.
' The following table suffices to show that in both cases infection
had taken place :—
(a) Piel Pier mussels.
No. 1. Contained 7 Cercarie.
No. 2. ¥ 3 live and 2 dead Cercariz, one of
* which was partly calcified.
No. 3. * 2 live and 2 dead Cercarie.
No. 4. 3A A live Cercariz.
No. 5. #5 2 live Cercarie.
(6) Specimens transferred from other tank at Brighton.
No. 6. Contained 19 Cercariz. :
No. 7. FF 3 Cercarie.
No. 8. Be 2 Cercarie.
No. 9 1 (dead) Cercaria.
Nor Tomer mminseesedl
The Adult Leucithodendrium,
Although I have not had an opportunity of making a direct
feeding experiment upon Somaterra or Gdemia, there is hardly any
doubt that the parasite that causes the formation of the pear-
sac, and consequently of the pearl, in Mytilus edulis is the larva
of Leucithodendrium somaterie (Levinsen), originally described
from the Hider Duck (Somateria mollissima Linn.) in Greenland,
158 DR. H, LYSTER JAMESON ON THE [Mar. 4,
and rediscovered by me in the Scoter, Wdemia nigra L., from
Bridlington Bay and the Villaine Estuary.
After finding the Sporocyst I made a careful examination of
such fishes and gulls as I could secure at Billiers, but could find
no parasite corresponding to the larva. However, on enquiring
of the fishermen I was informed that the great enemy of the
mussel in those waters is a diving duck, locally called ‘Cane
mouliére,” which frequents the Villaine in winter. M. d’Hamon-
ville, to whom I wrote on the matter, had no hesitation in saying
that this bird was @demia nigra. On wy visit to Billiers in
December last I proved, by shooting an example and procuring
four others that were taken in nets, that it was so. The name
‘Cane mouliére” seems to be applied to another duck as well,
probably the Scaup, Puligula marila Linn. A few young Scoters
remain on the Villaine during the summer months.
The Scoter is very common in winter at the mouth of the
Barrow Channel, just opposite the pearl-bearing mussel-beds.
Before going to Billiers in December I had proved the occur-
rence of LZ. somaterie (Levinsen), associated with Levinsenia
pygmeum Lev., ina specimen of Hdemia nigra sent me from
Bridlington by Mr. G. Williamson.
The Scoter received from Bridlington was in a rather advanced
state, and I could only determine the presence of a few examples
of LZ. somaterie in the hinder part of the small intestine. But
the five specimens procured on the Villaine were infested with
LL. somateric from the stomach to the anus and even in the ceca.
I caleulated that each specimen contained at least six thousand
examples of the parasite. Levinsenia pygmeum Linn., which
occurred abundantly in the Bridlington specimen, was not found
at Billiers. The intestine of the Bridlington duck contained
nothing but fragments of J/ytilus-shells. The example which I
shot at Billiers was feeding over mussel-beds, and the other four
were caught in special nets which are placed on the beds of
mussels at ebb tide, and in which the birds get entangled when
they visit these beds with the flood, to feed.
These Billiers specimens had apparently been feeding exclu-
sively on mussels, asno other food was found in them, except that
one individual contained about half a dozen Wucula sp. and a
broken Littorina-shell, in addition to Mytilus. In the crop and
stomach some of the mussels were still entire, and specimens up
to 40 millim. in length were found ; in the stomach the shells are
crushed, and pass through the intestine in small fragments at most
a few millimetres in diameter.
The striking likeness, except for the matter of size, between
the Mytilus-worm and Leucithodendrium somaterie, and the
occurrence of the latter in the two birds that are known to feed
par excellence on mussels, is almost sufficient to prove their
identity without the feeding experiment. I hope, however, to
make this experiment if birds can be secured. So far I have been
unable to purchase live examples, although I have made enquiries
1902. ] ORIGIN OF PEARLS. 159
in all directions. My discovery in the small intestine of a
Billiers Scoter, three inches behind Meckel’s diverticulum, of a
single immature example of the parasite, positively identical in
size and all details with the Cercaria from Mytilus, practically
proves the point.
The adults agree with Levinsen’s description and figure, except
that the genital pore (Pl. XVI. fig. 11, g.p.) is just in front,
and not in the centre, of the ventral sucker.
Levinsen’s observation on this point has been treated with
scepticism by later writers, and, indeed, such a position for the
opening of the penis and other genital tubes would not only be a
novelty in Trematode anatomy, but would probably render the
sucker useless as an adhesive organ.
The adult worm varies in size from *2 mm. to ‘55 mm, = It
is therefore only about half the size of the larva found in Mytilus.
It is protandrous, and specimens in which the male organs are
ripe are generally larger than egg-laden females.
The diminution in size accompanying sexual maturation is of
great interest, and can be accounted for in this way, I think.
The resting-stage in /ytilus is a highly assimilative phase in the
worm’s existence. The bulk of the body is greatly increased by
the distended gut and excretory system. A reduction in bulk
would be effected by the discharge of the contents of the latter,
but still more by the absorption of reserve material required to
mature the gonads. In fact, the reproductive organs seem to
grow at the expense of the other tissues of the body. The gut, I
may mention, is empty in the adult worm.
Specimens from (Wdemia die very quickly as a result of the
post-mortem cooling of the body of their host. The Scoter that I
shot was still warm when I dissected it, but the parasites died in
a few minutes when placed on a slide, and an hour after the bird
had been opened every adult worm was dead. The immature
specimen above mentioned survived the death of the host by
twelve hours. The temperature of the room where I worked at
Billiers was very low, and possibly in a well-heated laboratory
death would not ensue so quickly.
Owing to the rapid death of the worms, the material that I
preserved was not suited for detailed histological work, but the
relations of the various organs to one another could be determined
on sections, and by this means I have been able to check my
observations on pressure preparations of fresh material at Billiers.
Plate XVI. fig. 11 shows the structure of an average individual
in which the maximum number of eggs has not yet been reached,
and the uterus is not too complicated to mask the other structures,
The arrangement of the cuticular spines is the same as that
described for the larva. The suckers and the digestive and excre-
tory systems are also the same. The genital pore (g.p.) is just in
front of the ventral sucker. The penis (pe.) is pyriform. Its
extremity seems to be beset with glands. Posteriorly it goes over
into the large sac-like seminal vesicle (s.v.). This lies dorsal to
160 DR. H. LYSTER JAMESON ON THE [ Mar. 4
and, as a rule, a little to the right of the sucker. At its hinder
end it receives the vasa deferentia (v.d.). The testes (ée.) are a
little more comspicuous than in the larva. The uterus (wé.) opens
out just in front of the penis. The arrangement of its convolu-
tions depends upon the number of eggs contained (up to 100, or
even more). It seems to begin on the left side, near the ovary
(ov.), which is larger than the testes. It forms a series of con-
volutions on the left side, crosses in front of the genital opening
to the right, where it forms further convolutions, passes forward
as a loop to the anterior end of the body, and runs back to open
out at the genital aperture. I am not convinced that this arrange-
ment is always strictly adhered to.
The eggs in the uterus have the form depicted in my sketch
and in Levinsen’s figure. They measure from ‘018 mm. to
‘023 mm., the average dimensions being ‘021 mm. x ‘018 mm.,
but they differ considerably in different individuals. J can add
nothing new to Levinsen’s observations on the yolk-gland.
I sought in vain for this worm in Bernicla brenta Pall.,
Tadorna cornuta Gmel., Querquedula crecea Linn., Colymbus
arcticus Linn., Larus argentatus Gmel., and Rissa tridactyla
Linn. I have had no opportunity of examining other allied
birds.
Being unable to secure live Hiders or Scoters, I tried feeding a
male Pochard, Fuligula ferina Linn., with infected Mytili, but
without results.
L. somaterie will very probably be found in the other members
of the genera Somateria and Hdemia when looked for.
The Conditions essential to Pearl-production.
The characters and life-history of the parasite suffice to account
for the anomalous distribution of pearl-bearing mussels, and, by
analogy, throw light on the cause of the differences in the number
of pearls produced by the true Pearl-Oysters on various pearling
and shelling grounds.
In order to be abundantly infected Mytilus must be on the
bottom, for the tailless Cerearia or ‘“ Cercarieum” is dependent
upon its limited creeping powers, and the chance of being trans-
ported by currents and deposited with silt, &e.
Hence mussels grown on stakes, like those on Piel pier,
although right in the middle of the beds of pearl-hearing
individuals, are practically uninfected.
Secondly, there must be an abundance of the first host (Zapes
at Billiers, Cardiwm at Piel) in the immediate vicinity of the
mussels, in order to ensure frequent re-infection. For I find on
the coasts of the Villaine, where Tapes is scarce, the proportion of
infected Mytili is small. Moreover, on the Roosebeck Scar, outside
the Barrow Channel, where Cardium is not found, pearls do not
occur frequently.
Thirdly, the beds must be near the feeding-grounds of the
1902.] ORIGIN OF PEARLS. 161
Scoter (or another final host), and the set of the currents must
favour the transportation of the larvee or eggs (whichever it may
be) to the beds where 7'apes occurs.
Although it is only on certain beds that these conditions exist,
infection takes place to a small extent on very many beds. I have
hardly ever examined a sample of mussels from any locality with-
out finding here and there among them an infected individual.
Duration of Life of the Larva in Mytilus, and Rate of
Growth of Pearls.
Iam at present making experiments to test the longevity of
the resting larva. I have, however, three facts to record that
lead me to think it is less than two years.
While the mussels on the foreshore opposite Piel Fish-Hatchery
are highly infected, those on the Roosebeck Scar, outside the
Barrow Channel, are not so. When I was at Piel, Mr. Scott
showed me a small patch of mussels on the pearl-bearing beds, and
told me that these molluscs had been brought in from the Roose-
beck by a fisherman about two years previously and thrown down
there. JI examined a number of these mussels, and each of them
contained several small pearls. Some, indeed, had as many as
ten, and all were infested with the Trematode. From the presence
of pearls in these specimens, it is probable that the first Cercariz
to enter them had been dead some time. The dimensions of
these pearls throw some light on the time required to produce
pearls of a certain size. The five largest specimens weighed
together 6°9 mg. (dried on filter-paper after being preserved in
spirit), and measured respectively 1:3 x 1 mm., 1°55 x 1 mm.,
9 x ‘85 mm., 1:°2 x ‘8 mm., and 2:1 X 1:15 mm, The last was
obviously, from its form, a double pearl.
Again, as four out of the six specimens that I dissected after
they had been about two years in the Brighton Aquarium con-
tained each a small pearl, but no live Trematodes, it is probable
that the latter were unable to survive two years in the Aquarium.
Moreover, at Piel and Billiers pearls are very seldom found in
mussels less than 40 mm. long, which size is probably attained
when the mussel is in its third year. I find Cercariz, on the
other hand, in specimens only 20 mm. in diameter.
The average size of the larger pearls found in old mussels at
Piel is about 2x 2 mm., but all sizes, from the dimensions of the
parasite to 3°35 X 3°2 mm., were found. At Billiers they are
usually smaller, as the mussels are regularly fished there and
seldom reach a great age. The pearl-bearing beds at Piel are not
fished, as the infected mussels are not marketable.
The sizes to which pearls grow in other molluscs differ
very greatly for the several species and for the same species in
different localities. Their growth is, in fact, regulated by the
causes which control the thickening of the shell. Hence the
white porcellaneous pearls of Tridacna gigas and Hippopus
162 DR. H. LYSTER JAMESON ON THE [ Mar. 4,
hippopus, in which species the shell is thick and massive, are
often as large as a cherry. Among the true Pearl-Oysters the
large thick-shelled species, Margaritifera maxima Jameson and
M. margaritifera Linn., produce the largest pearls, those yielded
by the smaller J, vulgaris (Schumacher) on the Ceylon fisheries
and elsewhere being usually small, and prized rather for their
colour and lustre than for their dimensions.
And it is interesting to note that I. vulgaris in the Persian
Gulf, where it attains larger dimensions and yields a more massive
shell than in Ceylon, gives on an average larger pearls than the
Gulf of Manaar pearl-oyster.
The general experience of everybody acquainted with pearl-
fisheries is that the largest pearls are found in the oldest and
thickest shells, which proves how intimately the growth of pearl
and shell are associated. It is natural that such an association
should exist, since, as is obvious from the results recorded in this
paper, the mechanisms of both processes are the same.
Origin of Pearls in other Forms.
Two questions will naturally occur :—Are we warranted in
assuming that the mechanism of pearl-formation is the same in
other molluscs? and, Is it generally caused by Trematode larvee ?
Tn answer to the first question, I may say that in those cases
where I have been able to examine pearls in situ, in Margaritifera
margaritifera Linn., JM. vulgaris (Schumacher), J/. maaima
Jameson, Hippopus hippopus L., and Pinna nigrina Lam., this sac
is universally present. It has been noticed by von Hessling in
Margaritana margaritifera, and by Diguet in Margaritifera
margaritifera L., from California. But, apart from this evidence,
it is safe to say that without such an epithelial sac to shed the
cuticular conchyolin, the nacreous layers of the pearl could not be
laid down at all.
To what extent other causes besides Trematode larvee may be
capable of inducing such sacs to develop, has yet to be ascertained.
Trematodes have been unquestionably associated with pearl-
formation in Anodonta, Margaritana margaritifera, Mytilus
edulis, and Margaritifera (%) mazatlanica (see Introduction).
Besides these records, I have detected the remains of Trematodes
in decalcified or sectioned pearls from the following species :—
Margaritifera margaritifera Linn., M. maxima Jameson, WM. vul-
garis Schumacher, Pinna nigrina Lam., P. euglypia Hanley,
P. virgata Menke, Hippopus hippopus Linn., Tridacna gigas
Lam., and Vytilus magellanicus.
T examined pearls from several other molluscs, but had not
sufficient material to ascertain satisfactorily. In MW. vulgaris
Schumacher, besides the Trematode, there seems to be a second
organism, possibly a Gregarine, concerned in pearl-formation.
The periostracum pearls in the mantle-margin of Modiola are
also associated with parasitic protozoa.
1902. ] ORIGIN OF PEARLS. 163
These data suffice to show that in many molluscs, including
several of the species yielding the most valuable pearls, Trematodes
are one cause, if not the exclusive cause, of pearl-formation. To
what extent other parasites are capable of producing the same
effects cannot be said at present. That the other causes to which
pearls have from time to time been attributed play any part is a
matter of the merest conjecture only, and has never, so far as I
know, been demonstrated by experiment or investigation.
Possibilities of Economic Application.
The bearing of the facts recorded in this paper upon the
problem of artificially producing pearls, and so meeting the
difficulty presented by the increasing demand and exhausted
fisheries, is obvious. It was indeed with the hope of throwing
some light on this matter that I first took up the subject, about
three years ago. The key to the realizing of this, so often
regarded as an academic dream, lies obviously in the scientific
study of the parasites which occur in the valuable forms. This
was pointed out exactly fifty years ago by Filippi, but has been
ignored by most subsequent writers.
The life-history of the Trematodes occurring in the genus
Margaritifera probably agrees in the more essential points with
those of other Digenea. Their adult stages may reasonably be
expected to occur in the organisms that eat the pearl-oysters,
notably such fishes as Balistes, while the first host will almost
certainly be some mollusc occurring on the pearl-banks or shelling-
grounds.
Having ascertained the first host, there is no reason why
infection should not be performed by placing young pearl-oysters
in company with it in more easily accessible waters. To attempt
to establish the cultivation of pearl-oysters on new grounds
without also cultivating and infecting the first host of the
parasite would be futile. Needless to say, such methods of
artificially promoting natural infection would be incomparably
superior to any method of pearl-production by operation on the
individual oyster, as millions of examples could be treated by the
former method, while tens were being operated upon.
It is obvious from my Brighton experiment that infection can
be induced in Mytilus, and I can see no reason to doubt that, in a
couple of years, these MJyéili will contain pearls, resulting from
that artificially induced pathological condition.
LITERATURE.
1712. Mery.—Remarques faites sur la Moule des Estangs. Hist.
de Acad. Roy. des Sciences, Paris, Année 1710: Mém.
pp. 408-426.
1717. Réaumur.—Observations sur la Coquillage appellé Pinne
Marine ou Nacre de Perle. Mém. de ’Acad. Roy. des
Sciences, Paris, 1717, pp. 177-194, pls. v. & vi.
1858.
1858.
1858.
1859.
1859.
1871.
1872.
1877.
1881.
1882.
1885.
DR. H, LYSTER JAMESON ON THE [ Mar. 4,
. CHEMNITtz.—Vom Ursprung der Perlen. Der WNatur-
forscher, Bd. xxv. 8. 122. 1791.
. Homes, E.—On the Production and Formation of Pearls.
Phil. Trans. 1826, pt. 3, pp. 338-341, pl. xiii.
. D. C.—Some Account of the British Pearl-Fishery now
existing on the Conway. Loudon’s Mag. Nat. Hist.
vol. iil. pp. 1382—-134.,
. Dusarpin, F.—Histoire Naturelle des Helminthes ou Vers
intestinaux. Paris, 1845.
. Fiurprr, F. pe.—Sull’ origine delle Perle. Il Cimento,
fasc. iv. Torino, 1852.
. Fiureri, F. pe.—Encore un Mot sur la Formation des
Perles. Mill. Archiv, 1856, pp. 490-493.
. Frurprr, F. pe.—Troisieme Mém. pour servir 4 |’ Histoire
génétique des Trematodes. Mem. Accad. Torino, xviii.
pp. 201-232. 1859.
. Kécuenmetster.—Uber eine der hiufigsten Ursachen der
Elsterperlen. Miill. Archiv, 1856, p. 269.
. Kevaart, E. F.—Introductory Report on the Natural
History of the Pearl-Oyster of Ceylon. Proc. Phil. Soe.
Edinburgh, vol. i. pp. 399-405, 1854; also Madras
Journal, vol. ii, pp. 89-104, 1858.
Mésius, K.—Die echten Perlen: Hamburg, 1857. Abhandl.
Geb. Nat. naturw. Verein, iv. Bd. 1. Abth.: Hamburg,
1858.
von Huzsstinec.—Uber die Ursachen der Perlbildung bei
Unio margaritifer. Zeitschr. f. wiss. Zool. Bd. ix.
8. 543. 1858.
PacenstecHEeR, H. A.—Uber Perlenbildung. Zeitschr. f.
wiss. Zool. Bd. ix. 8. 496, Taf. xx. 1858.
Kevaart, E. F.—Report on the Nat. Hist. of the Pearl-
Oyster of Ceylon. Trincomalee, 1859.
Huxtey, T. H.—Tegumentary Organs. Todd’s Cyclopedia
of Anat. & Physiol. vol. v. pp. 490-1.
GarNER, R.—On the Formation of British Pearls and their
possible Improvement. Journ. Linn. Soc., Zool. vol. xi.
p. 426. 1871.
Hartine, P.— Recherches de Morphol. synthétique.
Amsterdam, 1872.
von NatrHusius-Koniesporn.— Untersuchungen tiber nich
cellulire Organismen. Berlin, 1877.
Levinsen, G. M. R.—Bidrag til Kundskab om Gronlands
Trematodfauna. K. D. Vidensk. Selsk. Oversigt, 1881,
pp. 52-84, tab. ii. & ii.
TuLuBerc, T.—Studien iiber den Bau u. das Wachsthum
des Hummerpanzer u. der Molluskenschalen. Sv. Ak,
Handl. Stockholm, xix. no. 3, pp. 1-57. 1882.
EurenspAum, E.—Untersuchungen iiber die Struktur u.
Bildung der Schale der in der Kieler Bucht haufig vor-
kommenden Muscheln. Zeitscher. f. wiss. Zool. Bd. xli,
S, 1-47, Taf. i. & ii, 1885,
1902. | ORIGIN OF PEARLS. 165
1885. Miuuer, Ferrx.—Uber die Schalenbildung bei Lamelli-
branchiaten. Schneider’s Zool. Beitrige, Bd. i. 8. 206-
246, Taf. xxvili._xxx. 1885.
1892. Moynier DE ViILLEPorx.—Recherches sur la Formation et
l Accroissement de la Coquille des Mollusques. Journ. de
VAnat. et de la Physiol. xxvii. pp. 461-518, pls. 19, 20.
1892.
1894. p’Hamonvitix, L.—Les Moules perliéres de Billiers. Bull.
Soc. Zool. France, 1894, pp. 140-142.
1894. Tuurston, E.—Pearl and Chank Fisheries of the Gulf of
Manaar. Madras Govt. Museum, Bull. No. 1. Madras,
1894.
1896. Loos.—Recherches sur la Faune parasitaire de Egypte,
Cairo, 1896.
1898. Compa, B.—La Madreperla. Torino, 1898.
1898. von Naruusius-Kénicspory, W.—Uber die Gestaltungs-
ursachen der Haare, der Eischalen, der Mollusken-
schalen und der Harting’schen Kérperchen. Archiv f.
Entwickelungsmechanik, vi. pp. 365-393. 1898.
1899. Srosstcu, M.— Lo smembramento dei Brachyccelium.
Trieste, 1899.
1899. Diever, L.—Sur la Formation de la Perle fine chez la
Meleagrina margaritifera. C.R. Acad. Sci. Paris,
exxvlil. pp. 1589-1591.
1901. Jameson, H. L.—On the Identity and Distribution of the
Mother-of-Pearl Oysters. P.Z.S. 1901, vol. i. pp. 372-
394.
1901. Brepermann, W.—Untersuchungen iiber Bau und Ent-
stehung der Molluskenschalen. Jenaische Zeitschr. f.
Naturwiss. xxxvi. pp. 1-164, Taf. i.—vi.
1901. Dusois, R.—Sur la Mécanisme de la Formation des Perles
fines dans le Mytilus edulis. C.R. Acad. Sci. Paris
cxxxlil. pp. 603-605.
EXPLANATION OF THE PLATES.
PuatTE XIV.
Mytilus edulis.
Fig. 1. Mytilus edulis L., Billiers. Section of a small Pearl, decalcified in situ,
showing remains of Trematode as nucleus. ewt.ep., external epidermis of
mantle; ¢.¢., connective tissue; b/., blood-corpuscles; s., epithelium of
pearl-sac; con., conchyolin basis of pearl; con.’, outermost uncalcified
layer of same, attached to epithelium ; cw., cuticle of dead Trematode; ph.,
pharynx, and dig., digestive system of same. X 90.
Fig. 2. The Trematode larva in the connective tissue of the mantle, prior to forma-
tion of sac. f,, fibres of connective tissue; p7., proliferating cells which
give rise to the epithelial sac; cw., cuticle; dig., digestive ceca; ew., excre-
tory organs, and pa., parenchyma of the parasite. Other figures as above.
130. The section passes between pharynx and ventral sucker. The
ventral surface of the parasite is turned towards external epidermis.
ig. 3. The cells of the proliferating epithelium which is destined to become the
sac (cf. fig. 12). mu., nuclei with chromatine reticulum ; ¢.t., connective
tissue.
Fig. 4. Cells of the fully-formed sac which surrounds the Cercaria in fig. 2. ¢.t.,
underlying connective-tissue cells and fibres.
a
r)
166 DR. P. L. SCLATER ON THE [ Mar. 4,
PLatE XV.
. 5. The sac fully developed around the larva, which is cut through ventral
sucker. s.v., rudimentary seminal vesicle at base of penis; c.m., circular
musculature ; l.m., longitudinal musculature; skz., ventral sucker. Other
letters as in previous figures. 180.
. 6. The Cercaria as it occurs in Mytilus. From a specimen stained in toto.
cu., cuticle ; m., mouth; a.s., anterior sucker; p.s., posterior sucker; ph.,
pharynx; s.gl., salivary glands; @., esophagus ; dig., digestive ceca; ex.,
excretory system; ex.p., pore of same; pe., penis; te., testes; v.d., vasa
deferentia. x 130.
Fig. 7. The same, in longitudinal section. pa., parenchyma; pa.n., nucleated sub-
cutaneous layer of same; int.ep., intestinal epithelium, in surface view ;
g-p-, genital pore; c.m., circular musculature; 1.m., longitudinal ditto.
m.f., muscle-fibre in parenchyma; 7.c., supra-cesophageal nerve-commissure
Other letters as in fig. 6. X 180.
Ki
gg
Fi
gg
Prats XVI.
Fig. 8. Dead Cercaria in Mytilus, with three centres of calcification. X 180.
Fig. 9. Section of the muscular mantle-margin of Tapes, showing the Sporocysts
with contained Cercarie. m., musculature of Tapes; sp., sporocyst ;
cer., Cercaria; ew., excretory tubes; dig., digestive ceca; pe., penis;
s., ventral sucker of same.
Vig. 10. Pressure preparation of the Cercaria from Sporocyst in Tapes. The speci-
men was examined alive, so sexual organs are not visible. e., eyes; t.p.,
tactile papille ; dig., digestive czeca; ea., excretory system. X 700.
Fig. 11. The adult worm from Cdemia nigra L., River Villaine. s.v., seminal
vesicle; ov., ovary; wt., uterus; g.p., genital pore. Other letters as in
fig. 6. Stained zm toto.
PratE XVII.
Fig. 12. Mytilus edulis L., Piel, Lancashire. Photo of a thin slice or “Schliff”
through centre of pearl, showing calcified Trematode as nucleus. X 25.
Fig. 13. Ditto. Pearl formed in sac vacated by Trematode. A few granules of
residual matter have calcified to form nucleus. X 265.
Fig. 14, Ditto, showing several centres of calcification in the Trematode. X 265.
Fig. 15. Ditto, double pearl, one of constituents being formed around Trematode,
the other as in fig. 18. X 25.
Fig. 16. Pearl from Margaritifera margaritifera Linn., New Guinea, showing
imperfectly calcified Trematode, with radially arranged prisms. X 25.
2. List of the Parrots represented in the Society’s Collection
in January 1902, with Remarks on some of the Rarer
Species. By P. L. Scnater, D.Sc, F.R.S., Secretary
to the Society.
[Received February 20, 1902.
(Plates XVIII. & XTX.,')
The birds in the Society’s Parrot-house having been lately
re-arranged and the duplicates and “ deposited” specimens having
been moved to another place, I have thought it worth while to
prepare a list of the species of Psittacide now represented in the
Collection, which may be useful in future years, and to add to it
a few notes on some of the rarer forms.
It may be observed that the 147 specimens of Parrots now
1 For explanation of the Plates, see p. 171.
DS IGOD sell Wl {Pil YOVAMII.
H.Goodchild del, MinternBros.Chnromo lith.
ECLECTUS WESTERMANNI,¢.
P.Z.5.1902 vol.I.Pl.xIx,
J-Smoxt del. et ith. Mantern Bros.Chromo:
PLATYCERCUS MASTERSIANUS.
¥
ae
a
—
=
. , i
Price 7 7
Was
=F) a
5 Pe!
, 4 |
- ’ \
bce
7 i
o-
‘ e
>
;, =
et]
i
=
‘
ik .
re
A ’
4
7.
im
ei
—
- 4 ~
oo
‘i
~
“ee
. , 5
ed
ti,
& *epgroe
eter
‘hs
=
a 2
' e
il
i). 3
é ; i
- re Five 6 iS
ine
1892. ] PARROTS IN THE SCCIETY’S GARDENS. 167
living in the Parrot-house are referable to the following 107
species *, which are arranged and named according to the order
followed in the last edition of the ‘ List of Vertebrated Animals,
(1896).
List or Livine Parrots, 1902.
Fam. NESTORID2.
*
—
. Nestor notabilis Gould.
Fam. LoRIip2&.
. Los rubra (Gin.).
riciniata (Bechst.).
wallacii (Finsch).
. Lorius domicella (Linvn.).
Trichoglossus hematodes (Linn.).
Sorstent (Temm.).
nove-hollandie (Giu.).
*9. —— rubritorques, Vig. & Horsf.
ornatus (Linn.).
11. Psitteuteles euteles (Temm.).
12. Glossopsitiacus concinnus (Shaw).
QO NI? OT YH Go bo
Fam. CACATUID2.
Subfam. CAacatuinz.
13. Calyptorhynchus naso Gould.
14. Callocephalon galeatum (Lath.).
15. Cacatua galerita (Lath.).
16. triton Temm.
Wie sulphurea (Gin.).
18. leadbeateri (Vig.).
16) moluccensis (Grn.).
#20. gymnopis Sclater.
21. sanguinea Gould.
22. ducorpsi Hombr. et Jacq.
23. hematuropygia (P. L. 8. Miill.).
24, roseicapilla VY ieill.
Subfam. CALOPSITTACINA.
25. Calopsittacus nove-hollandie (Gm.).
Fam. PSITTACcIDs.
Subfam. ConvuRINz
26. Anodorhynchus hyacinthinus (Lath.).
glaucus (Vieill.).
28. Ara ararauna (Linn.).
1 In March 1879 there were 170 Parrots in the Society’s Collection, referable to
98 species (see P. Z. 5. 1879, p. 299).
168
29. Ara macao (Linn.).
30. chloroptera Gray.
31. militaris (Linn.).
32. severa (Linn.).
33. maracana (Vieill.).
34. Conurus acuticaudatus (Vieill.).
35 auricapillus (Licht.).
36. —— nanday (Desm.).
37. —— rubrolarvatus Mass. et Souancé.
38. holochlorus Sclater.
39. —— ocularis Sclater & Salv.
40. —— cactorum (Max.).
Al eruginosus (Linn.).
42. Conuropsis carolinensis (Linn.).
43. Cyanolyseus patagonus (Vieill.).
44. Pyrrhura leucotis (Licht.).
45, Myopsittacus monachus (Bodd.).
46. Psittacula passerina (Linn.).
47. Brotogerys virescens (Gm.).
48, pyrrhopterus (Lath.).
49, twi (Gm.).
Subfam. Pronin a.
*50. Chrysotis guildingi (Vigors).
*O1, augusta (Vigors).
52. —— amazonica (Linn.).
53. cestiva (Linn.).
D4. ochrocephala (Gm.).
55. —— auripalliata (Less.).
56. levaillanti Gray.
57. ——- viridigena Cass.
58. salvimi Salvad.
59. —— autummalis (Linn.).
60. —— mornata Salvad.
61. —— festiva (Linn.).
*62. bouqueti (Bechst.).
63. albtfrons (Sparrm.).
64. ventralis (Miill.).
65. leucocephala (Linn.).
66. agilis (Linn.).
67. Pionws maximiliani (Kuhl).
68. chaleopterus (Fraser).
69. Caica melanocephala (Linn.).
DR. P. L. SCLATER ON THE
. Paocephalus gulielmi (Jard.).
Subfam. Psrrracinz
. Psittacus ertthacus Linn.
. Coracopsis vasa (Linn.).
nigra (Linn.).
[ Mar. 4,
1892. | PARROTS IN THE SOCIETY'S GARDENS.
Subfam., PALZoRNITHIN.
74, Hlectus pectoralis (Miill.).
0, westermannia Bp.
76. cardinalis (Bodd.).
17. Tanygnathus luzonensis (Linn.).
78. Paleornis torguata (Bodd.).
Co docilis (Vieill.).
80 Jfasciata (Miill.).
81 eupatria (Linn.).
82. magnirostris Ball.
83. —— derbianus Fraser.
84. —— schisticeps Hodgs.
*85 Jinschi Hume.
86. Polytelis barrabandi (Swains.).
87. melanura (Vig.).
88. Piistes erythropterus (Gm.),
89. Aprosmictus cyanopygius (Vieill.).
90. Pyrrhulopsis splendens (Peale).
91. Loriculus galgulus (Linn.).
92. vernalis (Sparrm.).
Subfam, PLATYCERCINA.
93. Platycercus elegans (Gm.).
94. Jlaveolus (Gould).
95), mastersianus Ramsay.
96. pallidiceps Vig.
97. cximius (Shaw).
98. —— barnardi (Lath.).
99. browni (Temm.).
100 semitorquatus (Quoy et Gaim.).
101. —— zonarius (Shaw).
102. Psephotus multicolor (Brown).
103. hematonotus Gould,
*104. chrysopterygius Gould.
105. Cyanorhamphus unicolor (Vig.).
106. novee-zealandie: (Gm..).
107. Welopsittacus undulatus (Shaw).
1. NESTOR NOTABILIS.
A pair of these birds were presented to us by the Hon. Walter
Rothschild, F.Z.S., on Feb. 16th, 1899.
169
They are fed upon
ordinary food, and have no meat given to them, as was, at one
time, thought to be necessary. They are in excellent health and
condition.
9, TRICHOGLOSSUS RUBRITORQUES.
Of this beautiful Australian species the four examples, deposited
by the Hon. W. Rothschild, F.Z.8., July 27th, 1900, are the first
Proc. Zoon. Soc,—1892, Vou. I. No. XI.
170 ON THE PARROTS IN THE SOCIETY’s GARDENS. _—[Mar. 4,
and only specimens ever received by the Society. (See P. Z.8.
1900, p. 772.)
20. CACGATUA GYMNOPIS.
This specimen is the oldest bird in the Parrot-house, having
been in the Society’s possession for 33 years. It was purchased
June 2nd, 1868.
50. CHRYSOTIS GUILDINGI.
51, 9 AUGUSTA.
62. - BOUQUETI.
We have always a good series of specimens of the Neotropical
genus Chrysotis in the Collection. We are now so fortunate as
to possess examples of these three rare Antillean species, and
only a short time ago had also an example of C-. versicolor of
Dominica, (See P.Z.S. 1890, p. 772.)
71. PsirraAcus ERITHACUS.
One of our specimens of this bird (received Dec. 24th, 1897)
has a pure white tail.
75. EcLECTUS WESTERMANNI. (Plate X VIII.)
In 1899 and 1900 we received under our care a series of ten
specimens of both sexes of this bird, deposited by Mr. Rothschild.
Whatever doubts may have been formerly expressed, it is now,
I think, quite certain that this is a valid species, although we do
not yet know its exact locality. Mr. Rothschild having already
published his notes on this remarkable species (Bull. B. O. C. x.
p. i, Oct. 1899), I need not repeat them here further than to
say that the male is at once recognizable by its entirely green
breast, and that the female is most like the corresponding sex of
L. pectoralis but has a dull purple (not blue) lower breast.
85. PALAORNIS FINSCHI.
This is a rare species from Burmah, of which we received our
first specimen in Nov. 1901 from Mr. E. W. Harper, F.Z.S., to
whom we are indebted for examples of so many rare Indian
species. (See Cat. B. xx. p. 458, pl. xii.)
95. PLATYCERCUS MASTERSIANUS. (Plate XIX.)
Platycercus mastersianus, Ramsay, Pr. Linn. Soc. N.S.W. i.
p. 27 (1877); Salvad. Cat. B. xx. p. 543.
We are so fortunate as to have a single example of this rare
bird in the Collection, deposited by Mr. Rothschild on the
29th Oct., 1897, and I have great pleasure in exhibiting a coloured
drawing of it prepared by Mr. Smit (Plate XIX.). It is clearly a
member of the group of P. elegans, but differs from all others
in having the central parts of the tail-feathers whitish. Count
Salvadori was unacquainted with it, and mer ely copied the original
description in a footnote.
1p AS OWA ol dle ILO.
W.Purkiss del.et lith. West ,Newman imp.
NEW SPECIES OF HALTICIDAS -
1902. ] ON NEW SPECIES OF AMERICAN COLEOPTERA. 171
104. PszPpHoTUS CHRYSOPTERYGIUS.
Of this species, perhaps the most beautiful of al] Australian
Parrakeets, we received a pair in immature plumage in March
1897. They are now in full plumage and in excellent condition.
EXPLANATION OF THE PLATES.
Prate XVIII. 8
Eelectus westermanni, @ and ? adult, p. 176.
PuatEe XIX.
Platycercus mastersianus, p. 170.
(Taken from specimen deposited on Oct, 29, 1897.)
3. Descriptions of New Species of Coleoptera of the Family
Haltiade from South and Central America. By
Martin J acosy, F.E.S.
{ Received February 15, 1902.]
(Plate XX.')
The constant application for the determination of so many
specimens of Halticide which J have received from different sources
has induced me to describe most of those which are contained in
my collection, for a long time unnamed, asa further contribution to
the extremely numerous described forms from Seuth and Central
America. The subject cam scarcely be dealt with at present im
anything but a very incomplete manner, but every little helps
and will one day assist in grouping together the enormous material
known, when the time and opportunity has arrived for a proper
monograph of this immense family, such as has been attempted
with the species known from Central America, in the great work
on that country by Godman and Salvin.
The present paper deals with that division of the Halticide m
which a more or less distinct thoracic sulcus is present in connection
with simple, not inflated, posterior claws.
DIPHAULACA COSTATIPENNIS, Sp. 1.
Dark metallic blue, the basal joints of the antenne fulvous;
thorax obscure cupreous, impunctate; elytra closely and nearly
irregularly punctured, the sides with two or three longitudinai
coste.
Length 4 millim.
Head impunctaie, the frontal elevations elongate, divided by a
deep groove, the carina acute; antennz long and slender, black,
the lower three joints fulvous below, stained with metallic blue
above ; thorax subquadrate, the sides straight at the base, rounded
1 for explanation of the Piate, seep. 204.
We
172 MR. MARTIN JACOBY ON NEW SPECIES [Mar. 4,
before the middle, the angles acute, the basal sulcus very deep,
straight, bounded at the sides by a perpendicular groove, the
surface entirely impunctate; elytra much wider at the base than
the thorax, the basal portion slightly raised, the dise very closely
and distinctly punctured, the punctures arranged in indistinct
rows, much finer towards the apex, a well-marked costa extends
from the shoulders nearly to the apex and is preceded by a shorter
one which does not extend upwards to the base, the interstices
between these costz more irregularly and more strongly punctured
than the rest of the surface, elytral epipleure broad, with a row
of punctures ; below and the legs metallic blue, impunctate, the
last segment of the abdomen with a longitudinal groove which
widens at the apex, the sides of the segment shghtly raised.
Hab. Venezuela.
The single specimen before me is. probably a male, although
the anterior tarsi are not particularly dilated. The species differs
from its allies in the sculpture and coste of the elytra, in connection
with its coloration.
DIPHAULACA FLAVIPES, Sp. Nn.
Below piceous, above greenish-eneous, the basal joints of the
antenne and the legs entirely flavous; thorax impunctate, with
deep transverse sulcus; elytra with the basal portion convex,
scarcely perceptibly punctured except near the base.
Length 4 millim.
Head impunctate, eneous, the frontal elevations small, ovate ;
antenne slender, blackish, the lower four joints flavous, the third
and fourth of equal length; thorax one-half broader than long,
the sides straight at the base, rounded anteriorly, the angles
acute, the disc deeply transversely sulcate, the sulcus sinuate,
bounded at the sides by a perpendicular groove, entirely im-
punctate; elytra widened towards the middle, the apex of each
rounded, the base distinctly swollen, the shoulders prominent,
the surface extremely finely punctured at the base, the punctures
searcely arranged in rows and very closely placed, nearly obsolete
below the middle, the epipleurz very broad, concave; below
obscure piceous, the legs entirely flavous.
Hab. Colombia (Pehlke). Museum Stettin and my collection.
This small species may be at once known by the entirely eneous
upper surface and the flavous legs. ‘The specimen before me seems
to be a female.
DIPHAULACA PALLIPES, Sp. Nn.
Below black, above greenish-zneous, the basal joints of the
antenne and the legs (the posterior femora excepted) flavous;
thorax impunctate, the basal sulcus deep ; elytra finely punctate-
striate, the punctures indistinct near the apex.
Length 2 millim.
Head impunctate, the frontal tubercles narrowly oblique, the
carina ucute, the lower portion of the face and the labrum black,
1902.] OF AMERICAN COLEOPTERA, 1¢3
the vertex greenish-eneous ; the antenne rather long and slender,
flavous, the terminal joints fuscous, the third and “fourth joints
equal, ‘scarcely longer but much thinner than the second one;
thorax twice as broad as long, the sides straight, the anterior
angles oblique, forming a distinct tooth before the middle, posterior
margin rather broadly produced at the middle, the basal sulcus
deep and sinuate, bounded at the sides by deep perpendicular
grooves, the surface rather convex, impunctate, greenish-eneous ;
elytra with the base slightly raised, distinctly punctured in closely
approached rows which become obsolete near the apex; legs
flavous, the posterior femora blackish.
Hab. St. Catharina, Brazil.
Of only half the size of D. Jlavipes, the basal margin of the
thorax not straight as in that species, but slightly produced at
the middle, the punctuation of the elytra more distinctly arranged
in rows, and the posterior femora black, not flavous.
DIPHAULACA FRUHSTORFERI, Sp. Nn.
Greenish-zneous, basal joints of the antenne and the base of
the femora (more or less) testaceous ; head deeply foveolate ; thorax
obsoletely punctured, metallic green or eneous, the basal sulcation
deep; elytra with the base raised, rather strongly and closely
punctate-striate, metallic green or brassy.
Length 33 millim.
Head impunctate, deeply foveolate between the eyes, the latter
large and prominent, frontal elevations obsolete, carina acute ;
antennz rather long and slender, piceous, the lower four joints
and the apical one testaceous, third and fourth joints equal,
terminal joints equally long, not thickened ; thorax transversely
subquadrate, the sides straight, the anterior angles slightly
thickened and oblique, the basal suleus deep, bounded at the sides
by equally deep perpendicular grooves, the surface convex, nearly
impunctate, brassy green; elytra with the base distinctly raised,
the basal margin preceded by a deep groove which extends in
front of the shoulders, the surface strongly and closely punctate-
striate anteriorly, the punctures very much finer towards the
apex, below dark eneous; legs robust, piceous, the femora more or
less testaceous at the base, posterior tibie slightly curved and
dilated at the apex.
Hab. St. Catharina, Brazil (Pruhstorfer).
Principally distinguished from other species of the genus by
the deep frontal fovea, the strong and close punctuation of the
elytra, and the colour of the legs.
DIPHAULACA HAROLDI, sp. n.
Below blackish, above greenish-eneous, the basal three joints
of the antenne flavous; thorax transverse, finely punctured ;
elytra with the basal por ‘tion convex, regularly. punctate- striate,
the strie remotely placed.
Length 23 millim.
174 MR. MARTIN JACOBY ON NEW SPECIES [| Mar. 4,
Head impunctate, the frontal elevations small, the carina
acutely raised; antenne extending to the middle of the elytra,
black, the lower three joints flavous, second and third joints equal,
fourth scarcely longer, the terminal joints gradually thickened ;
thorax quite one-half broader than long, the sides nearly straight,
the anterior angles obliquely thickened, the basal sulcus deep, not
extending beyond the perpendicular grooves at the sides, the
disc convex, remotely and finely punctured, elytra with the base
distinctly raised, punctured in regular and not closely placed.
rows, distinct to the apex, the last row near the lateral margins
more strongly punctured, the last interstice slightly convex ;
below and the legs nearly black; prosternum broadly elongate,
distinctly punctured.
Hab. Colombia.
One of the smallest species of the genus, and distinguished
from its Colombian congeners by the finely but distinctly
punctured thorax.
Genus Lacrica.
a. Hlytra blue or green.
LACTICA BOLIVIANA, Sp. 0.
Ferruginous, the antenne (the basal three joints excepted),
the apex of the tibie, the tarsi, and the abdomen black ; elytra
violaceous blue, strongly punctured ; thorax impunctate, the basal
sulcus deep; posterior legs entirely black.
Length 5 millim.
Head impunctate, obliquely grooved above the antenne, carina
short, labrum piceous; eyes very slightly emarginate; antenne
scarcely extending to the middle of the elytra, black, the lower
three joints fulvous, third and fourth joints equal, the second one
scarcely shorter; thorax twice as broad as long, rather convex,
not narrowed anteriorly, the sides straight, the anterior angles
oblique, the basal sulcus deep and straight, placed close to the
base, the surface impunctate ; scutellum fulvous; elytra convex,
violaceous blue, distinctly punctured in closely approached irregular
rows, distinct to the apex; legs and the breast ferruginous, the
apex of the four front tibie, the tarsi, the posterior legs entirely,
and the abdomen black.
Hab. Bolivia.
Closely ailied im coloration to LZ. elegantula Har., but much
larger, without distinct frontal elevations, the third and fourth
joints of the antenne equal; the head impunctate, and the elytral
punctuation distinct to the apex. JZ. abdominalis Jac. is much
smaller and has fulvous posterior legs.
LactTicA LOBATA, sp. n.
Oblong-ovate, ferruginous, apical joints of the antenne black ;
thorax impunctate, the basal sulcus rather deep, basal margin
1902. ] OF AMERICAN COLEOPTERA. 175
produced at middle; elytra metallic blue or greenish, finely
punctured in rows ; abdomen black.
Length 43 millim.
Head impunctate, the frontal tubercles entirely obsolete, the
carina short, pyriform; eyes large, nearly entire; antenne ex-
tending to about the middle of the elytra, black, the lower four
joints ferruginous, the third and fourth joints equal, but little
longer than the second joint; thorax twice as broad as long, the
sides very feebly rounded, narrowly marginate, the anterior angles
oblique, not produced, the basal margin rounded and produced at
the middle, the basal sulcus straight and moderately deep, the
perpendicular grooves very deep, the surface impunetate, ferru-
ginous; scutellum fulvous; elytra convex, subcylindrical, finely
punctured in closely approached rather regular rows, distinct to
the apex; breast and legs ferruginous; abdomen black.
fab. Bolivia.
In this species the basal margin of the thorax has a distinct
round lobe at the middle, differing in this respect from most of
the other species of the genus: this character alone would scarcely
justify the creation of another genus, but it will assist in the
recognition of the species, in connection with the system of
coloration and the black abdomen.
LacticA FUNEREA, sp. n.
Entirely dark violaceous, the last five joints of the antenne
fulvous; thorax transverse, impunctate, the basal sulcus deep ;
elytra purplish-violaceous, extremely finely punctured,
Length 6 millim.
Elongate, convex, the head impunctate, frontal elevations
entirely obsolete; eyes not very large; clypeus thickened, wedge-
shaped, carina narrow; antenne nearly reaching the end of the
elytra, black, the last five joints fulvous, the second half the
length of the third joint ; thorax strongly transverse, more than
twice as broad as long, the sides perfectly straight, narrowly
margined, the anterior angles oblique, the disc entirely impunctate,
blackish-violaceous, the basal sulcus and the perpendicular grooves
very deep; elytra metallic purplish-violaceous, very finely
punctured when seen under a strong lens; below and the legs
nearly black.
Hab. Yurimaguas, Peru.
Of this very distinct species, two specimens are in my
collection.
LACTICA COSTATIPENNIS, sp. n. (Plate XX. fig. 1.)
Flavous, the antennz (the basal joimts excepted) and the tibiz
and tarsi black; thorax impunctate, the basal sulcus deep ;
elytra bright green, finely rugosely punctate, the sides of each
with an acute costa.
Length 6 millim.,
Head flat and impunctate, the frontal elevations obsolete
176 MR. MARTIN JACOBY ON NEW SPECIES [ Mar. 4,
clypeus slightly raised, anterior edge of the labrum black; eyes
oblong, slightly sinuate; antenne extending to the middle of the
elytra, black, the lower three joints more or less flavous below,
the third joint smaller than the fourth, all the following joimts
elongate; thorax twice as broad as long, the sides rounded, the
anterior angles simple, not oblique, basal sulcus deep, slightly
sinuate, the surface flavous, impunctate; scutellum triangular,
fuscous ; elytra extremely closely and rather finely rugosely
punctured, not very shining, with an acute ridge from the shoulders
to the apex, accompanied by another shorter ridge below the
middle; below flavous, the lower portion of the tibie and the
tarsi fuscous.
Hiab. Llano Grande, Guatemala.
I have seen only a single example of this very distinct species,
which is in my collection.
LAcTICA BILINEATA, Sp. Nn.
Flavous, the antenne black; thorax transverse, impunctate,
with deep basal sulcus; elytra metallic blue, microscopically
punctured, each with a narrow flavous streak from the base to
below the middle.
Length 4 millim.
Head with a few deep punctures near the eyes, flavous, the
frontal elevations strongly raised ; clypeus flattened, subquadrate,
but slightly raised between the eyes; labrum piceous; eyes
very large, distinctly emarginate; antenne piceous, the second
joint half the length of the third, the other joints wanting ;
thorax more than twice as broad as long, very short, the sides
straight, the anterior angles oblique, the basal sulcus very deep,
placed close to the base and nearly straight, the dise impunctate ;
secutellum broad, black; elytra elongate and parallel, metallic
dark blue, with a few very fine punctures at the base, from the
middle of the latter proceeds a narrow flavous stripe which extends
to below the middle and is much constricted medially, its
direction is towards the suture; below stained with piceous, the
legs pale flavous.
Hab. Peru.
Easily recognizable on account of its coloration, which is
probably variable in regard to the elytral flavous stripes. I possess
only a single specimen of this very distinct species.
b. Elytra with spots or striped.
LLACTICA SEMINIGRA, sp. n. (Plate XX. fig. 2.)
Head and thorax flavous, the antenne, the breast, and legs
black; head and thorax, impunctate; elytra flavous, with a sub-
quadrate large black patch from the base to below the middle.
Length 6 millim.
Head impunctate, shining, flavous, with a small fovea between
the eyes, clypeus strongly raised in shape of a triangular ridge,
1902.] OF AMERICAN COLEOPTERA. at
antenne extending beyond the middle of the elytra, black, the
basal joint flavous below, the lower three joints shining, the rest
pubescent, third joint much shorter than the fourth; thorax
twice as broad as long, the sides straight, the anterior angles
oblique, the basal margin rounded and rather prominent at the
middle, the basal sulcus shallow, bounded laterally by deep per-
pendicular grooves, the surface impunctate; scutellum testaceous ;
elytra not perceptibly punctured, the anterior portion occupied by
a large black subquadrate patch which does not extend to the
lateral margins, the rest of the surface pale flavous, very shining;
the breast and legs black, the abdomen flavous.
Hab. Cachabé, Ecuador (Rosenberg).
I know only a single specimen of this very distinct species,
which is in my collection.
LACTICA BICOLORATA, sp. n. (Plate XX. fig. 3.)
Broadly ovate and convex, testaceous, the antenne, thorax,
legs, and the sides of the breast black ; elytra minutely punctured,
testaceous, the apical portion in shape of a transverse oblique
band, black.
Length 6 millim.
Head impunctate, the frontal tubercles flattened, testaceous ;
antenne extending to the middle of the elytra, black, the first
joint testaceous below, third and following joints elongate, rather
robust; thorax nearly three times broader than long, the sides
straight, the anterior angles oblique, the basal sulcus shallow, the
perpendicular grooves at the sides deep, the surface impunctate,
black; scutellum black; elytra widened posteriorly, minutely
punctured, testaceous, a transverse band, its inner margin concave
and widened at the suture, occupies about the apical fourth
portion of the elytra; below testaceous, the sides of the breast
and the legs black.
Hab. Peru.
Allied to ZL. batesi Baly, but with differently coloured antenne
and legs, and the black elytral portion narrower, the thoracic
sulcus also more shallow.
LAcTICA LIMBATIPENNIS, Sp. n.
Ovate, pale testaceous ; antenn very long ; thorax impunctate,
the basal suleus very shallow ; elytra extremely finely and closely
punctured, black, the lateral and apical margins narrowly pale
flavous.
Var. Elytra testaceous, the basal portion more or less black
only.
Length 2 millim.
Head impunctate, flavous; the eyes entire, frontal elevations
distinctly raised, clypeus flat; antenne quite extending to the
apex of the elytra, very slender, entirely flavous, the second and
following two joints gradually elongated ; thorax scarcely twice
as broad as long, the sides feebly rounded, the anterior angles not
178 MR. MARTIN JACOBY ON NEW SPECIES [ Mar. 4,
oblique, the basal sulcus very obsolete as well as the perpendicular
grooves, the dise impunctate, flavous; scutellum flavous; elytra
extremely closely and minutely punctured, convex, black, sur-
rounded by a pale flavous margin which widens rather at the
-apex ; below and the legs pale testaceous, the middle of the breast
sometimes stained with black.
Hab. Peru.
There are five specimens of this small species in my collection.
Of these two represent the variety which has the base of the
elytra black only and the rest of the dise faintly stained with
fuscous, so that the pale lateral margins are still plainly distinct.
c. Hlytra entirely pale-coloured.
LACTICA PARAGUAYENSIS, Sp. 0.
Flavous; antenne robust, black, the basal three joints flavous ;
thorax strongly transverse, the basal sulcus rather shallow ;
elytra parallel, flavous, impunctate, the apex of the femora
piceous, the tibiz and tarsi black.
Length 5 millim.
Of elongate, not very convex shape; the head impunctate with
the exception of a single deep puncture near the eyes, the latter
not large, frontal elevations only indicated, the carina very broad
and convex, clypeus likewise strongly raised ; labrum flavous, apex
of the mandibles and the palpi piceous; antennee comparatively
robust, black, the lower three joints flavous, stained with piceous
above, the second and third joints small, of equal length, the
following jomts rather dilated ; thorax more than twice as broad
as long, the sides straight, obliquely narrowed anteriorly, the
anterior angles obliquely produced, the basal sulcus shallow,
placed close to the base, the lateral perpendicular grooves very
deep, the surface entirely impunctate; scutellum rather broad,
triangular ; elytra impunctate.
Hab. Paraguay.
This Lactica, of which a single specimen is contained in my
collection, may be distinguished by the rather robust antenne,
the rather small-sized and widely separated eyes, and especially by
the very transversely shaped thorax and its produced and oblique
anterior angles. It differs in this respect from any of the many
similarly coloured species of the genus.
LACTICA APICIPES, sp. n.
Elongate, moderately convex, testaceous, the antenne (the
basal joint excepted), the apex of the tibiz, and the tarsi black;
thorax impunctate, transverse sulcus shallow, perpendicular grooves
deep; elytra extremely minutely punctured.
Length 4—5 millim.
Of elongate, parallel, and moderately convex shape, the head
impunctate, the frontal elevations obsolete, carina broad ; eyes very
1902. | GF AMERICAN COLEOPTERA. 179
large, slightly emarginate within ; antenne extending below the
middle of the elytra, black, the basal joint flavous, second joint less
than half the length of the third, the latter slightly shorter than
the fourth joint, terminal joints shorter than the intermediate
ones; thorax about one-half broader than long, rather flattened, the
sides very feebly rounded and very slightly narrowed anteriorly,
the anterior angles obsoletely oblique and slightly thickened, the
basal suleus moderately deep in the male, more shallow in the
female; elytra microscopically punctured; below and the legs
testaceous, the apex of the tibize and the tarsi black.
Hab. Balthazar (windward side), Grenada, W. Indies (Z. H.
Smith).
T identified this species formerly with Z. tibialis Oliv., but the
three specimens before me, after a closer examination, must
represent another species: they are of a very pale testaceous colour,
without punctuation of the head and thorax, and all have only
the apex of the tibize and the tarsi black, differing in that respect
from any of its allies. The specimen that I look upon as the male
is much smaller, the thoracic sulcus is deeper, and the anterior
tarsi have their first joint dilated.
LAcTICA WEISEI, sp. n.
Ovate, pale fulvous, the antennz (the basal joint excepted)
black, long and slender; thorax transverse, with deep basal sulcus,
impunctate ; elytra not perceptibly punctured; below and the legs
fulvous.
Length 4 millim.
Head with a transverse groove at the vertex and some deep
punctures near the eyes, the latter large, deeply emarginate, the
intermediate space with a very deep fovea, frontal elevations
absent; clypeus broad, ill-defined, and scarcely raised; antenne
nearly extending to the apex of the elytra, black, the lower three
joints fulvous, the third joint slightly longer than the second,
the following joints very elongate; thorax twice as broad as long,
the sides feebly rounded, with a distict narrow margin, anterior
angles obliquely thickened, the basal sulcus very deep and
stra'ght, the surface entirely impunctate; elytra ovate, im-
punctate; underside and legs fulvous; the last abdominal
segment triangularly emarginate. ( @ 2)
Hab. Pernambuco.
The structure of the head and the unicolorous legs well distin-
guish this species.
LACTICA STRIGATIPES, sp. n.
Flavous, the antenne (the basal joint excepted), the apex of
the femora above, and the tibiz and tarsi black; thoracic sulcus
deep; thorax and elytra impunctate.
Length 3} millim.
Of very convex shape, subcylindrical; the head impunctate,
with a rather deep impression above the eyes, the latter elongate,
180 MR. MARTIN JACOBY ON NEW SPECIES [Mar. 4,
slightly emarginate, frontal tubercles small and obsolete, labrum
piceous; antenne extending to the apex of the elytra, black, the
basal two joints flavous, stained with piceous above, third and
fourth joints equal; thorax transverse, slightly narrowed ante-
riorly, the sides nearly straight, the anterior angles strongly
oblique, the surface impunctate, the basal sulcus deep and
sinuate; elytra slightly widened towards the middle, impunctate ;
below and the femora flavous, the latter with a short black streak
above near the apex, the tibie and tarsi entirely black.
Hab. Pichindé, Colombia.
I cannot identify this species, of which three specimens are
before me, with any other described Zactica on account of the
long antenne, the entirely impunctate upper surface, and the
markings of the femora, which are the same in all the specimens.
There have been a great many species described of almost similar
coloration, of which Z. citrina Harold is perhaps the most closely
allied, but this species has a fine but distinct punctuation and is
without the femoral stripe.
LACTICA SEMIFULVA, Sp. n.
Rufous or flavous, the head and thorax black, shining and im-
punctate; elytra not perceptibly punctured; antenne and legs
flavous.
Length 4 millim.
Head impunctate, black, shining, the frontal tubercles obsolete,
the carina comparatively very broad and convex, clypeus deflexed,
labrum pale piceous; antenne long and slender, entirely fulvous,
the third joint slightly shorter than the fourth, this and the
following joints nearly equal; thorax transverse, about one-half
broader than long, the sides with a narrow margin, straight,
the anterior angles oblique, posterior acute; the disc impunctate,
black, shining, the basal sulcus deep, bounded at the sides by
another deep longitudinal groove; scutellum fulvous; elytra with
a shallow depression below the base, of a bright rufous or pale
fulvous colour, entirely impunctate, shining; the underside and
the legs flavous.
Hab. Espirito Santo, Brazil.
Of this very distinct species I possess three specimens, two of
which have the elytra of much paler colour than the other.
L. dichroa Har. seems to be a nearly allied species, but has the
antenne, the underside, and the legs black; this and v. Harold’s
species are the only members of the genus with which I am
acquainted having a black head and thorax and fulvous elytra.
LACTICA FLAVILABRIS, sp. 0.
Flavous, the head black; thorax short, strongly transverse,
impunctate; elytra elongate, parallel and strongly convex, rufous,
impunctate.
Length 6 millim.
Head black and shining, the eyes extremely large, the inter-
1902.] OF AMERICAN COLEOPTERA. 18]
mediate space very narrow, the frontal tubercles small, the carina
acute, the clypeus triangularly raised; labrum flavous, as well as
the palpi; antenn extending to the middle of the elytra, flavous,
the last two joints wanting, the second joint half the length
of the third; thorax twice as broad as long, narrowed to a
slight degree anteriorly, the sides straight, the anterior angles
oblique, the disc impunctate, flavous; the basal sulcus very deep,
sinuate, and placed rather closely near the basal margin ; scutellum
rather broad; elytra convex, elongate and widened towards the
middle, not perceptibly punctured ; below and the legs flavous.
Hab. Espirito Santo, Brazil.
Evidently allied to Z. capitata Illig., but rather smaller, more
convex, the antenne and the labrum entirely flavous, the first-
named shorter, the eyes still larger, and the elytra reddish-fulvous,
not testaceous.
LACTICA RUFO-BASALIS, Sp. Nn.
Elongate, parallel, flavous; the antenne (the basal joint
excepted) black; head and thorax reddish, impunctate, the basal
sulcus very deep; elytra impunctate, flavous, the base reddish
fulvous; below and the legs flavous.
Length 7 millim.
Head impunctate, reddish fulvous, very shining, the frontal
tubercles rather obsolete, carina acute and narrow; eyes large,
coarsely granulate, the intermediate space not broader than their
diameter; antenne extending beyond the middle of the elytra,
black, the basal joint flavous, the third joint twice as long as the
second, but shorter than the fourth joint; thorax about one-half
broader than long, slightly narrowed anteriorly, the sides straight,
with a rather broad, reflexed margin, the anterior angles oblique,
the disc extremely shining, impunctate, the basal sulcus very
deep, placed at some distance from the base, the lateral perpen-
dicular grooves still deeper; elytra subcylindrical, scarcely
widened at the middle, not distinctly punctured but crowded
with numerous piceous specks, the basal margin reddish, the rest
of the surface flavous, the metatarsus of the posterior legs as
long as the following joints together.
Hab. Tejuca, Brazil.
This is a large-sized species, of which a single example is in my
collection. It is distinguished by the coloration of the head,
thorax, and elytra, the flavous legs, and very deep thoracic
sulcus.
LActTICA FEMORATA, Sp. N.
Flavous, elytra rufous, the posterior femora piceous; head and
thorax impunctate, the basal suleus very deep and sinuate, elytra
impunctate.
Length 23 millim,
Head impunctate, eyes large, entire, frontal elevations obsolete,
carina short, acute; antenne entirely fulvous, extending beyond
182 MR. MARTIN JACOBY ON NEW SPECIES [ Mar. 4,
the middle of the elytra, the second and third jeints short, nearly
equal; thorax twice as broad as long, slightly narrowed anteriorly,
the sides nearly straight, the anterior angles oblique, the posterior
margin sinuate, the basal sulcus very deep and sinuate, the sur-
face impunctate, flavous; scutellum broad; elytra ovate and
convex, with a distinct depression within the shoulders, rufous,
shining and impunctate; underside and legs paler, the posterior
femora black except at the extreme base, metatarsus of the
posterior legs elongate.
Hab. Espix ito Santo, Brazil.
A small species, distinguished by the colour of the antenne, of
the elytra, and that of the posterior femora.
LACTICA CLARKI, sp. n.
Elongate, subdepressed, flavous; labrum and the antenne (the
basal two joints excepted), the knees, tibiw, and tarsi black ; thorax
not narrowed in front, impunctate, with deep basal sulcus; elytra
narrow and parallel, nearly impunctate.
Length 3 millm,
Head impunctate, the frontal tubercles broad, feebly raised, carina
likewise flattened and broad; labrum and palpi black; eyes large,
entire; antenne comparatively short and robust, black, the basal
two joints flavous, third joint scarcely shorter than the fourth;
thorax transverse, scarcely twice as broad as long, the sides
perfectly straight, not narrowed in front, the anterior angles
strongly oblique, the lower point of the angles in shape of a small
tooth, posterior angles likewise dentiform, the basal sulcus deep
and straight as well as the lateral perpendicular grooves, the
surface impunctate; scutellum broad, triangular; elytra with
some extremely minute punctures, only visible under a strong
lens; the knees, tibiew, and tarsi black, the rest of the underside
and the legs flavous.
Hab. Paraguay.
A distinct little species on account of the shape of the thorax,
which is not narrowed in front as in the majority of the species,
and the comparatively robust antenne; the general shape is also
less ovate and convex.
LAcTICA BREVICOLLIS, sp. n.
Elongate, moderately convex, testaceous; the antennz (the
basal joint excepted), the knees, tibie, and tarsi black; thorax
one-half broader than long, impunctate, the basal sulcus deep and
straight; elytra impunctate.
Length 43-5 millim.
Head impunctate, the frontal elevations distinct and rather
broad, the carina narrow and acute; eyes large, entire; antenne
extending beyond the middle of the elytra, black, the basal joint
flavous, third joint distinctly shorter than the fourth; thorax
only one-half broader than long, distinctly narrowed anteriorly,
the sides straight, with a very narrow margin, the anterior angles
1902.] OF AMERICAN COLEOPTERA. 183
but very slightly oblique, the basal sulcus very deep and straight
and placed at a proportionately long distance from the base, the
dise entirely impunctate; elytra wider at the base than the thorax,
not much convex and of parallel shape; below and the legs
flavous, the apex of all the femora and the tibie and tarsi black.
Hab. Colombia, Peru, Venezuela.
The comparatively narrow thorax, entirely impunctate upper
surface, the structure of the head (acute carina), and the black
apex of all the femora seem to distinguish this from any similarly
coloured species of Lactica with which I am acquainted; neither
can I identify the insect with L. bogotana Harold, of which but a
superficial description is given and which is described as 7 millim.
long, with closely approached eyes and a microscopic but distinct
punctuation.
LACTICA NIGRICORNIS, sp. n.
Broadly ovate, fulvous; the antenne, the knees above, the
tibie, and the tarsi black; thorax transverse, impunctate, the
basal suleus deep and straight; elytra minutely punctured in
semi-regular rows.
Length 5 millim.
Of more broadly ovate and less convex shape than the majority
of species, of a reddish-fulvous colour; the head impunctate, eyes
not very large, entire, frontal elevations entirely obsolete, clypeus
convex between the antenne ; labrum piceous, palpi black; the
antenne with rather short and robust joints, extending to about
the middle of the elytra, black, the basal joint more or less fulvous
below, second and third joints nearly equal; thorax twice as broad
as long, slightly narrowed anteriorly, the lateral margins perfectly
straight, the anterior angles oblique, not produced, surface im-
punctate, with a deep and straight basal suleus; elytra gradually
widened posteriorly, very finely punctured in irregular and closely
approached rows, indistinct at the apex; below and the legs paler,
the knees above, the tibiz, and tarsi black; tibiz clothed with
fine yellow pubescence.
Hab. St. Catharina, Brazil.
Much larger than ZL. tibialis Oliv., with entirely black antenna,
the elytra with a fine and distinct punctuation ; the general shape
resembling that of Haltica rufa Oliv. Two exactly similar speci-
mens are before me.
LACTICA GRACILICORNIS, sp. 0.
Oblong-ovate, fulvous; the antenne long and slender, black,
the lower three joints flavous; thorax transverse, impunctate, the
basal sulcus deep and sinuate, the sides anteriorly with a fovea;
elytra extremely finely and closely punctured.
Length 2 millim.
Head with a few punctures near the eyes, the latter large,
entire, frontal tubercles small and obsolete, carina narrow and
feebly raised, palpi flavous; antenne extending to the apex of
184 MR. MARTIN JACOBY ON NEW SPECIES Mar. 4,
r)
the elytra, black, the lower three joints flavous, basal joint
elongate, the second and third nearly equal, the latter, however,
much thinner, the following joints very elongate, the last three
thinner than the preceding joints; thorax rather more than twice
as broad as long, the sides very feebly rounded, the anterior angles
not oblique, the disc impunctate with a transverse impression at
the sides near the anterior angles, basal sulcus deep and sinuate,
closely placed to the basal margin; elytra convex, with a very
shallow depression below the base, extremely minutely and
irregularly punctured ; underside and the legs flavous.
Hab. Mexico.
Of this species I received two specimens from M. Deyrolle,
which had been found in Mexican tobacco. The small size, very
slender antenne, the anterior thoracic fovea, and the entirely
flavous legs will assist in its recognition.
Lactica CARINATA, Sp. 1.
Entirely testaceous, of elongate shape; head with a transverse
ridge; thorax impunctate, not very transverse, basal suleus deep
and straight; elytra not perceptibly punctured.
Length 44 millim.
Head impunctate; frontal elevations obsolete, replaced by a
transverse ridge between the eyes; the latter very large, occupying
the entire sides of the head; clypeus strongly and acutely raised
between the antenne; the latter extending beyond the middle of
the elytra, entirely testaceous, the third joint one-half longer
than the second, but much shorter than the fourth joint; thorax
searcely twice as broad as long, slightly narrowed anteriorly, the
sides nearly straight, with a narrow reflexed margin, the anterior
angles oblique, but not produced, the basal sulcus and the
perpendicular grooves deep, the disc impunctate; elytra nearly
parallel, impunctate ; below and the legs coloured like the upper
parts, the metatarsus of the posterior legs elongate.
Hab, Cayenne.
There will be no difficulty in distinguishing this species from
its allies on account of the transverse ridge between the eyes and
the unicolorous legs. I know only a single specimen of this
species, which is in my collection.
LACTICA IMPRESSICOLLIS, sp. n.
Ovate, fulvous, the antennz (the lower three joints excepted)
black ; thorax transverse, with an antemedial transverse depression
at the sides, the basal sulcus deep and sinuate; elytra with a
shallow depression below the base, impunctate ; legs fulvous.
Length 4 millim.
Head deeply punctured near the eyes, the latter large, strongly
emarginate, the intra-ocular space raised in two elevations ;
clypeus bluntly elevated ; antenne extending beyond the middle
of the elytra, black, the lower three joints flavous, the second one-
half shorter than the third joint, the others closely pubescent ;
1902. ] OF AMERICAN COLEOPTERA. 185
thorax twice as broad as long, the sides feebly rounded near the
base, very slightly narrowed anteriorly, the anterior angles
oblique, the basal sulcus deep and rather sinuate, the anterior
portion of the thorax impressed with a transverse fovea at the
sides, the surface impunctate; elytra widened towards the middle,
feebly transversely depressed below the base, impunctate, their
epipleurz very broad and concave.
Hab. Bahia.
Amongst the unicolorous species, the present Lactica is dis-
tinguished by the raised intra-ocular space in shape of two blunt
elevations, and by the antemedian fovea of the thorax.
LAcTICA BAHIAENSIS, Sp. n.
Narrowly oblong, entirely testaceous; thorax impunctate, the
basal sulcus deep and very sinuate; elytra not perceptibly
punctured.
Length 4 millim.
Head with a deeply punctured sulcus round the eyes, the latter
large, feebly emarginate, frontal elevations only indicated ; clypeus
strongly convex between the antenne ; the latter extending below
the middle of the elytra, entirely testaceous, the second joint one-
half shorter than the third, the fourth longer than the preceding
joint; thorax about twice broader than long, the sides straight,
the anterior angles oblique, the basal sulcus deep and strongly
bisinuate, the disc impunctate; elytra not perceptibly punctured.
Hab. Bahia.
I must separate this species from its unicolorous allies on
account of the more than usually sinuate thoracic sulcus, in
connection with the structure of the head.
Lactina Harold.
LACTINA LEVICOLLIS, sp. Nn.
Dark blue, the thorax nearly subquadrate, entirely impunctate ;
elytra extremely closely and subrugosely punctured, clothed with
very short grey pubescence.
Length 6-7 millim.
Male. Head impunctate, the frontal elevations strongly raised,
elongate and oblique, carina acute, labrum black; antenne nearly
extending to the apex of the elytra, dark blue, the third and
fourth joints equal; thorax about one-half broader than long, the
sides nearly straight, the anterior angles slightly produced out-
wards, the base with the usual deep, sinuate transverse sulcus, the
surface entirely impunctate; elytra widened near the apex,
extremely closely and rather finely punctured, the interstices very
finely wrinkled and clothed with very short greyish hairs; epi-
pleure broad, concave and glabrous; the male organ curved and
like that of Z. glabrata, but with the anterior cavity shorter and
more ovate, the sides raised and with a blunt ridge within.
Hab. Peru.
Proc. Zoou. Soc.—1902, Vou. I. No, XITI. 13
186 MR, MARTIN JACOBY ON NEW SPECIES [ Mar. 4, -
I must separate this species from any other on account of the
impunctate thorax, which even under the strongest lens shows no
trace of any punctuation. Examples of both sexes are before me,
and do not seem to vary except in the slightly broader thorax
of the female.
LACTINA SEMIRUGOSA, Sp. n.
Metallic green, glabrous above, below bluish; thorax sub-
quadrate, finely punctured, the transverse sulcus deep; elytra
widened below the middle, very closely and strongly punctured,
the interstices more or less rugose.
Length 7 millim.
Head very finely transversely wrinkled, without punctures, the
frontal elevations strongly raised, pyriform, carina acute ; antenne
long and slender, metallic blue at the four or five basal joints,
the rest black, third and fourth joints equal, shorter than the
fifth ; thorax about one-half broader than long, the sides nearly
straight, very slightly narrowed anteriorly, the surface very
finely but not very closely punctured, the basal sulcus deep and
nearly straight, bounded laterally by a deep perpendicular groove ;
scutellum blackish; elytra widened below the middle, without
basal depression, comparatively strongly and very closely punctured,
with finely wrinkled interstices ; below very sparingly pubescent.
Hab. Colombia.
I know of no other species of Zactina with such strongly
punctured elytra in connection with the glabrous upper surface.
The description is that of the male, in which the anterior tarsi
are dilated, and the last abdominal segment has a central narrow
groove; the female does not differ except in the more slender
tarsi and the generally rather larger size.
LACTINA GLABRATA, Sp. N.
Metallic dark blue, glabrous above; thorax one-half broader
than long (¢), nearly impunctate, with deep sinuate sulcus ;
elytra very closely and distinctly punctured.
Length 6 millim.
Male. Head impunctate, the frontal tubercles strongly raised,
oblique, the carina acute; the antenne nearly extending to the
apex of the elytra, metallic blue, the third joint slightly longer
than the fourth and as long as the fifth joint; thorax one-half
broader than long, the sides straight at the base, slightly rounded
anteriorly, the anterior angles thickened, the surface with a deep
sinuate sulcus, the anterior portion rather convex, very minutely
and somewhat closely punctured as well as the base below the
sulcus; scutellum small, impunctate; elytra convex, not widened
posteriorly, attaining their greatest convexity at the middle, very
closely and rather strongly punctured, the apex of each rounded,
their epipleure concave, impubescent; below and the legs smooth,
shining ; the male organ curved, parallel, the apex rather blunt,
with an elongate cavity.
1902. ] OF AMERICAN COLEOPTERA, 187
Hab. Colombia, Venezuela.
I know of only one other described species which has an
impubescent upper surface (L. chalcoptera Har.); but in that
species the carina of the head is short and blunt, the thorax is
impunctate, the elytra have traces of coste and are cupreous in
colour. In the female of the present species the antennz are
much shorter and the elytra show a slight depression below the
base; the insect is also much more robust in shape.
DisonycHA DECEMMACULATA, Sp. 0.
Black; thorax flavous, with four black spots; elytra strongly
punctured, black, each with five flavous spots (2, 2, 1), the one
near the scutellum curved; the femora and the abdomen flavous.
Length 4 millim.
Head with a deep punctured groove near the eyes, black, with
a flavous mark on the vertex, the frontal elevations distinct, the
clypeus strongly convex, flavous ; labrum black ; antennz robust,
black, the lower four joints flavous below, third joint slightly
shorter than the fourth; thorax twice as broad as long, flavous,
the sides straight, the anterior angles scarcely oblique, rather
rounded, posterior angles oblique, the disc impunctate, with four
small black spots placed transversely ; elytra strongly and closely
punctured, black, each with five yellow spots, one at the humeral
callus, another in shape of a 0 near the scutellum, two round
spots at the middle and a transverse one near the apex; the
breast, the tibize and tarsi black; femora and abdomen flavous,
the apex of the posterior femora black.
Hab. Pernambuco, Pery-Pery.
This species will be easily recognized by the strong elytral
punctuation and the outwardly curved spot near the scutellum,
DIsONYCHA ELONGATA, Sp. 0.
Elongate and subdepressed, testaceous, the terminal joints of
the antenne fuscous; thorax impunctate; elytra not perceptibly
punctured ; a very narrow sutural and a sublateral stripe and a
broader longitudinal band at the dise, black.
Length 9 millim.
Head impunctate, the clypeus with an acutely raised central
ridge ; eyes reniform, rather deeply emarginate; antenne extend-
ing to the middle of the elytra, rather robust, fuscous, the lower
four joints testaceous, the fourth joint nearly twice as long as
the third; thorax about one-half broader than long, the sides
nearly straight with a narrow margin, the posterior angles
strongly oblique, anterior angles obliquely truncate; scutellum
black; elytra rather flattened, not perceptibly punctured, tes-
taceous, very shining, the suture very narrowly and a slightly
wider stripe close to the margins black, another much broader
band extends from the middle of the base nearly to the apex;
below and the legs testaceous, the apex of the posterior tibis
obscure fuscous.
13*
188 MR. MARTIN JACOBY ON NEW SPECIES | Mar. 4
Hab. Venezuela.
This species closely resembles in coloration and pattern many
others of the genus, but may be distinguished by the impunctate
elytra, the width of their black stripes, the comparatively narrow
and unspotted thorax, and the rather large general size of the
insect.
DiIsoNYCHA BREVICOLLIS, sp. n.
Oblong, pale testaceous, the antenne (the basal joints excepted)
and the tarsi black; thorax short and transverse, scarcely
punctured ; elytra closely and distinctly punctured, the sutural
and lateral margins and a narrow obsolete discoidal stripe obscure
pale fulvous.
Length 6 millim.
Head with a few punctures placed transversely between the
eyes, the frontal elevations feebly raised; clypeus broad, tes-
taceous, like the labrum and the rest of the head; eyes rather
elongate; antennz robust, black, the basal three joints flavous ;
thorax more than twice as broad as long, the sides nearly straight,
the anterior angles produced obliquely outwards, the posterior
angles oblique but not produced, the basal margin rounded, the
dis® with a few fine punctures; elytra closely and comparatively
strongly punctured, the sutural and discoidal stripes very narrow
and obsolete, the lateral margins more broadly marked with pale
fulvous; below testaceous, the tarsi black.
Hab. Ventanas, Mexico.
This species, which was unknown to me during the publication
of the Mexican Phytophaga in the ‘ Biologia Centr.-Amer.,’ is
readily distinguished by the shortness of the thorax, the more than
usual strong elytral punctuation, and the narrowness and faint
coloration of their stripes. ‘Two specimens are in my collection.
DIsoNYCHA ANGULATO-FASCIATA, sp. n. (Plate XX. fig. 4.)
Flavous, the head with two, the thorax with five black spots ;
elytra impunctate, black, a transverse medially constricted band
at the base, another -at the middle, and a spot near the apex
flavous; the apex of the femora and the tibie and tarsi
black.
Length 4 millim.
Head with some deep punctures near the eyes, flavous, a spot on
the vertex and another small spot between the eyes black, the
latter rather large; carina strongly raised; antenne robust,
black, the lower four joints flavous, the basal two stained with
black above, third and fourth joints equal; thorax twice as broad
as long, the sides straight, very narrowly margined, the anterior .
angles oblique, the disc impunctate, flavous, with five small black
spots (2, 3) placed transversely, the outer ones of the second row
the largest, the base with an obsolete transverse sulcus ; scutellum
black ; elytra not perceptibly punctured, with alternate transverse
flavous and black bands of angulate shape, the sutural and lateral
1902.] OF AMERICAN COLEOPTERA. 189
margins likewise black; below flavous, the anterior legs entirely,
the apex of the femora and the tibie and tarsi black.
Hab. Pernambuco, Serra de Communaty.
This little species resembles somewhat the well-known D. austri-
aca Schauf., but the design of the elytra is different and consists
of two flavous and two black transverse angulate bands and a
flavous spot at the apex: the first of these light bands is placed
somewhat obliquely at the base, leaving the humeral callus black,
the second is situated at the middle and is strongly constricted
medially.
CACOSCELIS GUIANAENBIS, Sp. Nn.
Flavous, the vertex of the head, the antennz, the femora above,
and the tibiz and tarsi black ; thorax finely punctured, with two
black spots ; elytra metallic green, strongly and closely punctured,
the lateral margins narrowly flavous; tibiz not emarginate.
Length 10-12 millim.
Hab. British Guiana.
I am obliged to separate this species from the well-known
C. marginata Fab., which it completely resembles in coloration, on
account of the entire tibiz, the same parts in C. marginata having
a very distinct emargination. Other less striking differencesgire
to be found in the rather stouter antenne of the present species,
in the more transversely shaped thorax, which is closely and
finely punctured. at the sides and has two large well-separated
blackish spots, with sometimes a smaller intermediate one.
Fabricius’s species is generally found in Brazil. In the present
insect the vertex of the head is likewise black, which I have
not found to be the case in any specimen of the allied species.
C. compta Erichs. has the sutural margins flavous as well as the
lateral ones. ‘There are four specimens before me.
CACOSCELIS TIBIALIS, Sp. 0.
Below flavous, above fulvous, the antenne, tibie, and tarsi
black or piceous; thorax transversely subquadrate, impunctate,
obsoletely sulcate; elytra finely and closely punctate-striate.
Length 7-8 millim.
Of elongate and nearly parallel shape, the head impunctate,
fulvous, the frontal elevations transverse, bounded behind by a
deep groove; clypeus rather strongly raised between the antenne,
its anterior margin straight ; labrum flavous, with a few punctures ;
terminal joint of the palpi acutely pointed; antenne extending
slightly beyond the middle of the elytra, black, slender, the third
and fourth joints more slender than the following ones but not
longer, basal joint more or less fulvous below ; thorax about one-
half broader than long, of equal width, the sides rounded, the
anterior angles somewhat oblique, the lateral margins accompanied
by a depression, the base with a shallow transverse groove; the
disc impunctate, fulvous, shining; scutellum triangular, im-
punctate; elytra finely punctured in closely approached, rather
190 "MR. MARTIN JACOBY ON NEW SPECIES [Mar. 4,
regular rows, their epipleure very broad, continued to the apex ;
below flavous, the tibie and tarsi black (sometimes only infuscate),
the posterior tibie with a small spur, the others unarmed; pro-
sternum narrow, anterior cavities open.
Hab, Espirito Santo, Brazil.
What I take to be the female of this species differs in having
a rather broad reflexed lateral margin to the elytra, and the upper
surface of a more pronounced fulvous; otherwise there is no
difference of any importance. The species differs from C. flava
Clark and C. testacea Cl. in the general smaller size, the arrange-
ment of the elytral punctuation, and in the black tibiz and tarsi.
In the female the elytral interstices are more or less costiform.
The species, like several others placed in this genus, has entire,
not emarginate tibie.
CACOSCELIS VARIPES, Sp. 0.
Below black; head and thorax fulvous, impunctate; elytra
dark greenish, opaque, minutely punctured; the anterior and
intermediate femora fulvous, the tibie and tarsi and the posterior
femora black.
Length 10 millim.
Head extremely minutely punctured when seen under a strong
lens, fulvous, the frontal elevations subquadrate, rather strongly
raised; clypeus with a highly raised blunt ridge between the
antenn, the last-named organs black, the basal joint fulvous, the
terminal two joints very elongate, much longer than the preceding
ones; thorax nearly twice as broad as long, slightly narrowed
anteriorly, the sides feebly rounded, with a rather broad flattened
margin, the base with a shallow sulcus, not extending to the
sides, the anterior angles slightly thickened, the surface im-
punctate, fulvous; scutellum broad, black; elytra slightly wider
at the base than the thorax, rather convex, of opaque greenish
colour, very finely punctured, with a narrow reflexed lateral
margin, their epipleure very broad and concave; below clothed
with grey pubescence, black, the anterior and intermediate femora
fulvous; tibiee not emarginate at the apex, the first joint of the
posterior tarsi as long as the following joints together.
Hab. Brazil.
Of this species, which differs in coloration from any of its
allies, a single specimen is in my collection, without any exact
locality. It is somewhat allied in colour to C. opacipenmis Jac.,
from Colombia, but differs entirely in the shape of the thorax
and the colour of the underside and legs.
CACOSCELIS CARULEIPENNIS, Sp. 0.
Pale fulvous, the antenne (the first joint excepted) black; head
and thorax impunctate; elytra dark violaceous blue, finely and
closely punctured, with traces of longitudinal sulci.
Length 7 millim.
Head fulvous, entirely impunctate, the frontal elevations -
1902.] OF AMERICAN COLEOPTERA. 191
distinct, eyes oblong; antenne extending beyond the middle of
the elytra, black, the basal joint fulvous, the third and following
two joints elongate, equal; thorax strongly transverse, slightly
narrowed anteriorly and at the base, the sides rounded anteriorly,
with a very narrow margin, the surface impunctate, light fulvous ;
scutellum of similar colour; elytra very distinctly, closely, and
somewhat rugosely punctured, with some very obsolete longi-
tudinal sulci, dark violaceous blue; below and the legs pale
fulvous, the tarsi infuscate.
Hab. Brazil.
Of this species I know only a single specimen, which is in my
collection. It is allied to C. violaceipennis Clark, but the elytra
are without the flavous lateral margins, the tibie are fulvous, not
black, and the antennz have the basal joint fulvous only.
OcNOSCELIS BOLIVIANA, sp. n.
Testaceous ; antenne greenish black; head and thorax greenish
or fulvous. nearly impunctate; elytra closely and distinctly
punctured, testaceous or obscure fulvous, the base broadly and the
sides narrowly metallic green.
Length 5 millim.
Male. Of ovate and depressed shape, the head strongly punctured
at the vertex, more or less metallic greenish, the frontal tubercles
very strongly developed, pyriform; antenne as long as the body,
blackish, the basal joint strongly thickened, the third and fourth
equal, terminal joints very elongate and slender; thorax com-
paratively long, scarcely one-half broader than long, the sides but
feebly rounded, with a narrow flattened margin, the posterior
angles acute, the anterior angles obsoletely thickened, the disc
nearly impunctate and smooth, more or less metallic green, with
the sides narrowly fulvous or entirely of the latter colour ;
scutellum blackish ; elytra closely and rather strongly punctured,
obscure fulvous, the base more or less metallic green, this colour
also extending down at the sides to about the middle of the
elytra; below and the legs testaceous, the tibiz and tarsi gene-
rally darker, the intermediate tibiz of the male strongly curved.
Hab. Bolivia.
Like the other three species of this genus, the present one
seems rather variable in regard to coloration, but in seven
specimens before me the colour of the elytra is constant. The
shape of the thorax and the impunctate disc of the latter distin-
guish the species from its allies. In the female the antennz
are shorter, the thorax is more transverse and is finely punctured.
Nepurica Harold.
This genus has been established by von Harold on a species
having the general appearance of a Disonycha or Asphera, but in
which the eyes are reniform and emarginate, the sides of the
thorax with a narrow margin, and the tibie without any emargi-
192 MR, MARTIN JACOBY ON NEW SPECIES [Mar. 4,
nation near the apex. All these characters are, however, not
wanting in many species of Disonycha, and intermediate stages
frequently occur in which it is impossible to say to which genus
to refer the species. Vephrica is therefore not a well-founded
genus, and it is impossible to fix the limit between this and
Disonycha. Nevertheless I have left the question in its present
state, and described all those species contained in my collection
in which the kidney-shaped eyes and other details peculiar to
von Harold’s genus are well marked, so that there will be little
difficulty in recognizing these forms.
NEPHRICA BOLIVIANA, Sp. N.
Pale flavous, the base of the head, the antenne, underside, and
legs black; thorax impunctate; elytra extremely minutely
punctured, yellowish white, a broad transverse band at the base
and another below the middle, not extending to the lateral nor
apical margins, metallic green.
Length 6 millim.
Head piceous at the base, impunctate, the lower portion nearly
white, the carina distinct and broad; antenne not extending to
the middle of the elytra, piceous, the basal joint flavous below ;
thorax twice as broad as long, the sides very feebly rounded, with
a narrow margin, the anterior angles truncate, the posterior
slightly truncate, the surface impunctate, yellowish white; scu-
tellum black; elytra nearly impunctate or with some extremely
minute punctures at the sides, metallic green, this colour not
quite extending to the lateral nor apical margins, and divided at
the middle by a narrow transverse flavous band; below and the
legs black (in immature specimens stained with flavous).
Hab. Bolivia.
Allied to WV. didyma Mllig. and WV. kirschi Har., but differing in
the design and colour of the elytra. In the above given description
I have taken the green colour for that of the ground, but in the
diagnosis the pale colour.
NEPHRICA MACULIPENNIS, sp. n. (Plate XX. fig. 8.)
Below black, as well as the antenne and legs ; head and thorax
flavous, the inten with five small black spots (4, 1); elytra closely
punctured, testaceous, the basal margin, the shoulders, a narrow
lateral stripe, connected with a broad transverse band below the
middle, and a round spot near the latter part, metallic green.
Length 7 millim.
Head sparingly punctured near the eyes, flavous, the extreme
vertex black; caria acute, labrum and palpi black; antenne
short, not extending ‘to the middle of the elytra, Sled the lower
three joints testaceous below, third joint shorter than the fourth,
terminal joints slightly thickened ; thorax twice as broad as long,
the sides feebly rounded, the anterior angles oblique, the surface
impunctate, flavous, with four small piceous spots placed trans-
versely anteriorly, and another spot at the middle near the base ;
1902. ] OF AMERICAN COLEOPTERA. - 1938
scutellum black; elytra very closely and strongly punctured,
testaceous, with the basal and sutural margins, a humeral spot,
and a narrow submarginal stripe metallic green ; this latter stripe
joins a transverse broad band before the apex, while a similarly
coloured spot is placed at the middle of the disc on each elytron;
below and the legs black, finely pubescent.
Hab. Rio Grande do Sul, Brazil.
NEPHRICA SANGUINOLENTA, sp.n. (Plate XX. fig. 10.)
Below fuscous, above dark red; thorax obsoletely sulcate, im-
punctate; elytra very minutely punctured, bright red, the disc
with a large subquadrate yellowish patch.
Length 8 millim.
Head impunctate, with the exception of a single puncture near
the eyes, the latter kidney-shaped, large; clypeus thickened, but
rather broad between the antenne; the latter extending to the
middle of the elytra, dark fulvous, the third joint twice as long
as the second and as long as the fourth; thorax rather more
than twice as broad as long, the sides nearly straight, slightly
obliquely narrowed towards the apex, with a distinct reflexed
margin, anterior angles thickened, slightly produced outwards,
the disc impunctate, very obsoletely suleate near the base, dark
reddish; elytra closely and extremely minutely punctured, of
nearly blood-red colour, the latter interrupted at the middle by a
large subquadrate yellowish patch, the anterior edge of which
extends rather nearer towards the base than the posterior one
towards the apex ; below and the legs dark fulvous.
Hab. Espirito Santo, Brazil.
A single specimen of this well-marked species is contained in
my collection.
NEPHRICA FULVICORNIS, sp. 0.
Flavous, the antennz and the legs fulvous, the breast and the
posterior femora black; head and thorax impunctate; elytra
bluish black, a transverse band at the middle, the apex and the
lateral margins flavous.
Length 5 millim.
Head impunctate, pale flavous; eyes large, sinuate, frontal
tubercles distinct ; clypeus slightly raised, triangular ; antenne
robust, fulvous, the third joint distinctly shorter than the fourth ;
thorax more than twice as broad as long, the sides with a rather
broad flattened margin, anterior angles obtusely thickened,
the disc impunctate, yellowish white; scutellum black; elytra
perceptibly punctured on the pale portion only, bluish black, this
colour interrupted by a transverse flavous band at the middle
which does not quite extend to the suture, but is connected with
the similarly coloured lateral margin, the apex likewise pale
flavous ; below flavous, the sides of the breast and the posterior
femora black.
Hab. Amazonia.
194 MR. MARTIN JACOBY ON NEW SPECIES [ Mar. 4,
This species differs from 1. brasthensis and WV. boliviana in the
flavous apex of the elytra and in its general smaller size, also
in the colour of the underside and legs.
NEPHRICA PARAGUAYENSIS, Sp. 1.
Dark fulvous; thorax with rounded sides, impunctate ; elytra
very closely and finely punctured, flavous, the basal and sutural
margin, a narrow transverse band before and another below the
middle, dark fulvous.
Length 6 millim.
Head finely punctured between the eyes, the frontal elevations
obsolete, the carina broad ; antennz fulvous, the third and fourth
joints equal; thorax twice as broad as long, the sides strongly
rounded and broadly margined, the anterior angles obtusely
rounded, the surface impunctate, fulvous ; elytra closely and finely
punctured, flavous, the base with a narrow fulvous band extend-
ing to the shoulders, two other medially constricted bands, one
before, the other below the middle, extend across the elytra but
do not reach the lateral margins ; below and the legs dark fulvous.
Hab. Paraguay.
Two exactly similar specimens are before me, which resemble
greatly in their markings certain species of the genus Homopheta.
NEPHRICA INCLUSA, sp. n. (Plate XX. fig. 5.)
Yellowish white, the antenne, tibie, and tarsi black; thorax
impunctate, the sides straight; elytra extremely finely punctured,
the basal and sutural margins, a transverse band before the middle
enclosing a basal spot of the ground-colour, and another trans-
verse band before the apex, reddish fulvous.
Length 7 millim.
Head yellowish white, impunctate, the vertex with a black
band; the frontal tubercles, the labrum, and palpi black; clypeus
very broad; antenne black, the third and fourth joints equal ;
thorax with the sides nearly straight, the anterior angles broadly
truncate, the surface impunctate, yellowish white; scutellum
black ; elytra extremely finely and rather closely punctured, pale
flavous, the base and a transverse band before the middle, con-
nected at the shoulders by a narrow longitudinal stripe, fulvous,
another narrow band below the middle and the suture narrowly
of the latter colour; below and the legs flavous, the posterior
femora above and the tibi and tarsi black.
Hab. 2
This species, of which I unfortunately do not know the locality,
but which is probably from Brazil, much resembles WV. para-
guayensis, but has nearly white upper and under sides, and the
thorax is of totally different shape, having straight instead of
rounded sides, the elytral punctuation also is much finer.
NEPHRICA CLAVERI, sp. n. (Plate XX. fig. 9.)
Head, the underside, and the posterior femora black; the
1902.] OF AMERICAN COLEOPTERA. 195
antenne, thorax, and the legs flavous ; thorax impunctate ; elytra
extremely finely punctured, black, a transverse band before and
another one below the middle flavous.
Length 6 millim.
Head black, with a few punctures near the eyes, the frontal
tubercles and the carina acutely raised; antenne extending to
the middle of the elytra, flavous, the basal three joints stained
with black above; thorax of the usual shape, the sides feebly
rounded at the middle, the surface impunctate, flavous, the base
with a rather well-marked transverse sinuate sulcus; scutellum
black ; elytra extremely minutely punctured, black, with a slightly
curved flavous transverse band at the middle and another near
the apex, their epipleure, the underside, and the posterior femora
black; the anterior legs, the posterior tibie, and all the tarsi
flavous; tibiz entire, not emarginate.
Hab. Colombia, Ibague (Prére Claver).
Of this very distinct species I received a specimen from Frére
Sebastien at St. Génis Laval, which was obtained by Frére Claver,
an ardent explorer of parts of Colombia.
NEPHRICA STAUDINGERI, sp. n. (Plate XX. fig. 11.)
Flavous, the base of the head, the antenne and legs fulvous;
thorax impunctate, flavous; elytra dark fulvous, impunctate, a
round spot at the base, a transverse band at the middle, another
near the apex, and the lateral margins narrowly, flavous.
Length 5 millim.
Head with one or ¢wo deep punctures near the eyes, the vertex
piceous, the lower portion flavous, frontal elevations obsolete, the
carina distinct; labrum piceous; the antenne dark fulvous, the
fourth joint slightly longer than the third ; thorax with the sides
nearly straight, the anterior angles truncate but not produced,
the base with an obsolete transverse sulcus, the surface impunctate,
flavous ; scutellum flavous; elytra entirely impunctate, dark
fulvous, each elytron with a round flavous spot at the base, a
slightly curved transverse band at the middle, and another one
near the apex, the lateral margins likewise very narrowly flavous
as well as the outer margin of the elytral epipleure; below
flavous, the legs dark fulvous.
Hab, Amazonia.
This little species very much resembles Disonycha austriaca
Schauf., but the elytra are fulvous, not black, and have flavous
lateral margins.
NEPHRICA TERMINATA, sp. 1.
Black, the thorax testaceous, impunctate, the disc black ; elytra
very finely and closely punctured, testaceous, the basal margin
and a triangular apical spot black.
Length 53 millim.
Head impunctate, with the exception of a deep puncture near.
the eyes, black, shining ; the base of the antenne and the clypeus:
196 MR. MARTIN JACOBY ON NEW SPECIES | Mar. 4,
flavous, labrum and palpi black; antenne rather slender, black,
all the joints, with the exception of the second, nearly equal ;
thorax with the sides straight, the anterior angles broadly truncate
and rather produced, the surface with the usual obsolete trans-
verse sulcus near the base, impunctate, testaceous, the dise with
a triangular black band or spot at the middle; scutellum black ;
elytra with the base narrowly black, this colour extending to the
shoulders, the apex with another triangular black spot ; below and
the legs black, the base of all the femora flavous.
Hab. Upper Amazons.
NEPHRICA BRASILIENSIS, sp. n. (Plate XX. fig. 6.)
Pale fulvous, the head, antenne, and the legs black; thorax
yellowish white, impunctate, scutellum black; elytra scarcely
perceptibly punctured, black, shining, the lateral margins and a
large transverse patch at the middle yellowish white.
Length 8 millim.
Head black, with a single puncture near the eyes, the latter
very large, kidney-shaped; clypeus flavous, triangularly raised,
labrum and palpi black; antenne rather short and robust, black,
the third joint twice as long as the second and nearly equal to
the fourth joint, intermediate joints slightly widened; thorax
nearly three times broader than long, slightly narrowed in front,
the sides nearly straight, with rather broad, strongly reflexed
lateral margins; anterior angles obtusely thickened, the surface
impunctate, very shining, nearly white, with a very obsolete
sulcus near the base; elytra convex, widened towards the middle,
with narrow lateral margins, nearly impunctate, shining black,
this colour interrupted at the middle by a large transverse
yellowish patch, which does not quite extend to the suture, but is
connected at the sides with the similarly coloured lateral margin ;
below pale fulvous, the legs black, the extreme base of the
posterior femora fulvous.
Hab. Rio Janeiro.
A rather large and convex species, well distinguished by its
coloration.
NEPHRICA IMITANS, sp. 0.
Testaceous, the antenne, tibie, and tarsi black; thorax im-
punctate, obsoletely sulcate near the base; elytra impunctate, a
broad sutural and discoidal band and a very narrow sublateral
stripe black.
Length 7 millim.
Head impunctate, the frontal tubercles obsolete, the carina
acutely raised, eyes broadly emarginate ; antennz black, the basal
two joints testaceous below, the fourth joint longer than the
third; thorax of usual transverse shape, the sides very feebly
rounded at the middle, the anterior angles strongly obliquely
truncate, the posterior margin emarginate in front of the scu-
1902.] OF AMERICAN COLEOPTERA. 197
tellum ; the dise impunctate, testaceous, very obsoletely trans-
versely suleate near the base; scutellum black, margined with
testaceous; elytra rather broadly ovate, impunctate, with a broad
sutural and a very narrow sublateral black band, the dise with
another broad band not quite extending to the apex, their epi-
pleure testaceous, black at their inner portion near the base;
below and the legs testaceous, the tibiz and tarsi blackish.
Hab, Peru.
Shorter and broader than Disonycha elongata, the sutural band
much wider and the antenne and legs differently coloured ; the
thorax is also more transverse in shape and the posterior angles
less obliquely cut; the elytral pattern resembles entirely that of
many species of Disonycha, but the eyes are distinctly reniform
and the thorax is of different shape.
NEPHRICA NIGROFASCIATA, Sp. Nn.
Black, the apical joints of the antenne and the legs flavous ;
thorax impunctate, flavous; elytra very closely and distinctly
punctured, black, a subsutural and a sublateral narrow band,
connected at the apex, flavous, apex of the posterior femora black,
Length 8 millim.
Head black, shining, impunctate, with the exception of a few
punctures near the eyes, the latter very large, reniform ; antenne
with the basal and the last four joints flavous, the others black;
thorax twice as broad as long, the sides straight, the angles
obliquely truncate, the surface flavous, impunctate; elytra closely,
strongly, and irregularly punctured, flavous, the sutural and
lateral margins narrowly black, the dise occupied by a broad
longitudinal band which does not extend to the apex; below
black, the femora and tibiz flavous, the apex of the posterior
femora and all the tarsi black.
Hab. Espirito Santo, Brazil.
A large species, easy of recognition on account of the colour of
the antenne and strong elytral punctuation, and resembling much
a species of Disonycha, from which the shape of the eyes will
distinguish it ; the basal margin of the thorax is scarcely oblique
at the posterior angles and altogether different in shape than in
Disonycha.
NEPHRICA UNIFASCIATA, sp.n. (Plate XX. fig. 7.)
Black, the thorax fulvous, impunctate, obsoletely sulcate ;
elytra black, shining and impunctate, a round spot at the base
of each and a transverse band near the apex white; abdomen
fulvous.
Length 6 millim.
Head black, with a single deep puncture near the eyes, the
latter large, reniform, frontal tubercles obsolete, carina strongly
raised ; antenne black, the second and third joints small,
nearly equal, the fourth elongate; thorax scarcely twice as broad
198 MR, MARTIN JACOBY ON NEW SPECIES [Mar. 4,
as long, the sides straight, not narrowed anteriorly, with a very
narrow margin, anterior angles feebly truncate, the surface
impunctate, fulvous, with an obsolete shallow suleus near the
base, posterior angles scarcely oblique; scutellum black; elytra
very shining, black, slightly depressed below the base, not
perceptibly punctured, a round spot, placed at the middle of the
base, and a transverse, very regular band near the apex yellowish
white ; below and the legs black; abdomen fulvous.
Hab. Peru.
A very distinct species on account of its coloration.
SYSTENA CLARKI, sp. 0.
Testaceous ; thorax transversely subquadrate, impunctate ;
elytra dark fuscous, the basal margin narrowly testaceous, the
surface impunctate.
Length 3 millim.
Head impunctate, testaceous or pale fulvous, the frontal
elevations distinct, subquadrate; antennz entirely flavous, the
third joint not longer than the fourth, the terminal joints slightly
thickened; thorax about one-half broader than long, the sides
rounded at the middle, distinctly constricted at the base, the
angles acute, the surface with a shallow but distinct basal sulcus,
impunctate, obscure testaceous or fulvous; scutellum flavous;
elytra wider at the base than the thorax, entirely impunctate,
dark fuscous or nearly black, the base narrowly testaceous; below
and the legs testaceous; prosternum extremely narrow, the
posterior femora but moderately thickened, with a piceous short
streak above near the apex, the metatarsus of the posterior legs
elongate.
Hab. Colombia.
There are three specimens of this species contained in my
collection ; the thorax is of rather convex shape, more so than is
generally the case in this genus, and the first joint of the posterior
tibize is proportionately long, but there is nothing to separate the
species generically.
SYSTENA PUNCTATISSIMA, Sp. N.
Below black, above pale testaceous, antennz fuscous; the head
remotely, the thorax and the elytra very closely and finely
punctured.
Length 4 millim.
Head distinctly but not closely punctured, the frontal tubercles
feeble, the labrum piceous; antennz rather stout, entirely pale
fulvous, the joints slightly stained with fuscous above, terminal
joints slightly stouter and shorter than the preceding ones;
thorax one-half broader than long, the sides distinctly rounded
at the middle, the anterior angles rather obsolete, the surface
very closely punctured at the sides, less closely at the middle,
the basal suleus very feeble; elytra with a slight transverse
1902. ] OF AMERICAN COLEOPTERA. 199
depression below the base, punctured like the thorax and of the
same colour, the suture slightly infuscate; below black, legs
testaceous, the posterior tibie deeply channelled at the apical
portion and also emarginate.
Hab. La Plata.
Of this very distinct species I possess two specimens ; in one of
these the basal joint of the antenne and the apex of the others is
more distinctly marked with fuscous.
SYSTENA BRASILIENSIS, sp. n.
Testaceous, the antenne (the basal joints excepted). black ;
thorax very minutely punctured, the sides with or without a
black stripe ; elytra extremely finely and closely punctured, black,
the lateral and apical margins narrowly testaceous; abdomen
black.
Length 5 millim.
Head impunctate, testaceous or flavous, the frontal elevations
feebly raised, clypeus rather depressed anteriorly, scarcely separated
from the face ; antenne not extending to the middle of the elytra,
black, the lower four joints testaceous below, the terminal five
joints distinctly shorter than the preceding ones; thorax of
usual shape, the sides straight at the base, slightly rounded
anteriorly, the anterior angles obtuse, the basal sulcus shallow
but distinct, slightly sinuate, the surface with a few scarcely
perceptible punctures, testaceous, the sides with a narrow black
band; scutellum black; elytra with an obsolete depression below
the base, very closely and finely punctured, black, the lateral
margins narrowly and the apical one more broadly pale testaceous ;
below and the legs testaceous, the abdomen black, the posterior
femora black at the apex.
Hab. St. Catharina, Espirito Santo, Brazil.
There are six specimens of this distinct species before me, in
some of which the lateral thoracic stripe is more or less or
entirely obliterated. The species may be distinguished from
similarly coloured varieties of S. variabilis Jac., from Panama, by
the black abdomen, the shorter antenne, the more distinctly
marked thoracic suleus, and the black apex of the posterior
femora.
SYSTENA ABBREVIATA, Sp. 0.
Below black, basal joints of the antennz fulvous, lower part of
the face flavous; thorax finely punctured, testaceous, with a
transverse black band; elytra minutely punctured, testaceous, a
subsutural and a sublateral stripe, abbreviated behind, black; legs
testaceous.
Length 5 millim.
Head very finely wrinkled at the vertex, the latter piceous or
black, the lower portion flavous ; antennz piceous, the lower four
joints flavous, fourth joint longer than the third ; thorax twice
as broad as long, the sides feebly rounded before the middle,
200 MR. MARTIN JACOBY ON NEW SPECIES [ Mar. 4,
narrowly emarginate, the anterior angles obtuse, posterior angles
distinct, the base with a narrow transverse sulcus, the disc
scarcely perceptibly punctured and wrinkled, black, all the margins
narrowly flavous; scutellum black; elytra sculptured like the
thorax, the fine punctures closely placed, the ground-colour
flavous or testaceous, a rather broad longitudinal stripe near the
suture, and another narrower one near the lateral margins, both
abbreviated behind, black.
Hab. Puebla, Mexico.
Of this species, which seems to be very rare, as I did not meet
with it in the numerous collections I had before me when working
out the Central American fauna, I have two exactly similar
specimens now: they differ from the other species in the black
thoracic band, the two posteriorly abbreviated elytral stripes, and
the black underside ; of the stripes, the subsutural one is of the
same width as that of the following flavous space, but the lateral
one is narrower. JD. discicollis Clark has a black head, differently
coloured legs and elytra, the latter are black with a discoidal
flavous stripe.
OXYGONA BRASILIENSIS, sp. n.
Testaceous or pale flavous, the antennz piceous; thorax im-
punctate, not strongly transverse, the angles acute; elytra finely
and closely punctured.
Length 6 millim.
Head broad, impunctate, without any fovez, the frontal
elevations strongly raised; the clypeus triangular; the antennze
piceous or fuscous, nearly extending to the end of the elytra,
the third and following joints elongate, equal, terminal joints
shorter ; thorax not much more than one-half broader than long,
the sides rather strongly rounded anteriorly, all the angles acute,
the anterior ones slightly oblique but scarcely produced, the surface
smooth and shining, with a narrow margin at all the sides; scu-
tellum more or less fuscous; elytra extremely closely and rather
finely punctured, the apex nearly impunctate ; below and the legs
testaceous, the metatarsus of the posterior legs elongate, the pro-
sternum extremely narrow, the last abdominal segment of the
male triangularly emarginate at the apex, with a narrow central
groove.
Hab. Espirito Santo, Brazil.
I must separate this species from 0. acutangula Chev., on
account of the much longer and much less transversely shaped
thorax, which is very obvious when the two insects are compared
in both sexes. O. luridulus Cl. and O. simplex Cl. have both
flavous antenne; the former has also a medial fovea on the head,
but what Clark meant bya short, deep, and broad medial marking
on the head, of which he says nothing in his description, it is
difficult to understand. Two specimens of the present species are
in my collection,
«
1902.] - OF AMERICAN COLEOPTERA. , 201
OXYGONA NIGRICOLLIS, sp. n.
Black, head and thorax impunctate ; elytra pale flavous, nearly
impunctate.
Length 5-6 millim.
Head entirely impunctate, with an oblique depression above
the eyes, the frontal tubercles strongly raised, the carina short
and broad; antenne black, extending beyond the middle of the
elytra, the third and the following two joints very elongate,
the others shorter; thorax more than twice as broad as long, the
sides subangulately rounded before the middle, strongly narrowed
at the base, with a broad reflexed margin, the anterior margin
accompanied by a narrow sulcus, the disc entirely impunctate,
shining, black; scutellum black; elytra pale flavous, extremely
minutely punctured ; below and the legs black,
Hab. Espirito Santo, Brazil.
This typical Oxygona, of which three specimens are before me,
is well distinguished by its system of coloration.
CREPIDODERA FLAVOMACULATA, Sp. Nl.
Piceous, above black, the basal joints of the antenne and the
legs (the posterior femora excepted) testaceous ; thorax opaque,
minutely punctured and granulate, feebly transversely sulcate ;
elytra very closely punctate-striate, piceous, the humeral callus
and the apex flavous.
Length 22 millim.
Head entirely impunctate, black, the frontal elevations obsolete,
clypeus convex between the antennz, labrum flavous; antenne
extending beyond the middle of the elytra, testaceous, the
terminal joints more or less darkened, second and third joints of
equal length, shorter than the fourth, terminal joints thickened ;
thorax subquadrate, one-half broader than long, the sides nearly
straight, the angles distinct, the surface opaque, finely punctured
and minutely granulate, the basal sulcus feeble, not extending
to the sides, the space below it also distinctly punctured ;
scutellum black, impunctate; elytra wider at the base than the
thorax, extremely closely punctured in irregular rows, the apex
much more finely punctate, the disc blackish, the shoulders with
a small flavous spot, the apex more or less broadly of the latter
colour, the space along the suture somewhat depressed; below
piceous, the legs testaceous or flavous, the posterior femora piceous,
their metatarsus as long as the two following joints together, all
the tibiee with a small spine, prosternum very narrow ; the anterior
coxal cavities closed.
- Hab. Concepcion, Talcahuano, Chili.
The thorax in this species is of an opaque, silky appearance
and the sulcus but feebly impressed, although distinct. I cannot
refer this insect to any of the species described by Philippi from
the same country.
Proc. Zoou, Soc.—1902, Vou. I. No. XIV. 14
202 MR. MARTIN JACOBY ON NEW SPECIES [ Mar. 4,
CREPIDODERA (CHALCOIDES ¢) ERICHSONI, Sp. n.
Greenish-zeneous, above metallic greenish-cupreous, antennz
flavous ; thorax convex, finely and closely punctured, the basal
suleus indistinct, the lateral grooves deep; elytra with deep
basal depression, distinctly and regularly punctate-striate, tibize
more or less fulvous.
Length 3 millim.
Of rather broadly ovate and convex shape, the head impunctate,
with the exception of a few fine punctures near the eyes, the
latter with oblique grooves from the inner margin to the middle
of the head, the frontal tubercles obsolete, the carina strongly
raised, terminal joint of the palpi acute and slender ; the antenne
rather long, flavous, the third joint very slightly shorter than
the second, the fourth and following joints more elongate,
terminal ones slightly thickened; thorax transversely convex,
the sides nearly straight, the anterior angles oblique, slightly
thickened, the basal margin sinuate near the scutellum, rather
broadly produced at the middle, the basal sulcus feeble at the
middle, more deeply impressed at the sides, slightly sinuate,
limited laterally by deep, somewhat curved, perpendicular grooves,
the dise strongly convex, closely and finely punctured; scutellum
small, blackish ; elytra scarcely wider at the base than the thorax,
the basal portion rather strongly raised, bounded by a transverse
depression, the disc convex, subcylindrical, finely and regularly
punctate-striate, metallic green with coppery reflections ; below
greenish-eneous, distinctly punctured, the legs piceous, the tibiz
obscure fulvous at the base, tarsi blackish, the first joint of the
posterior tarsi as long as the following two joints together ; pro-
sternum rather broad ; coxal cavities closed.
Hab. Peru.
The single specimen of this species contained in my collection
does not quite agree with any of the genera allied to Crepidodera,
on accountof the produced median lobe of the thorax, which gives
the latter quite a different appearance from that of the other
species; this lobe is not pointed but strongly rounded, and the
species ought perhaps to be placed in a special genus, but this may
be deferred until more similarly structured species turn up.
CREPIDODERA MAGISTRALIS, Sp. Nn.
Rufous, the antenne (the basal joints excepted) and the tibiee
and tarsi black; thorax transverse, impunctate, with deep basal
sulcus; elytra metallic greenish, very closely semi-punctate-striate.
Length 7 millim.
Head impunctate, frontal tubercles acute, carina short; antennz
extending to about the middle of the elytra, black, the lower
three joints fulvous, the third joimt shorter than the fourth ;
thorax about two-thirds as broad as long, the sides rounded, the
anterior angles slightly thickened and produced, the basal sulcus
deep and slightly sinuate, bounded at the sides by a perpendicular
1902.] OF AMERICAN COLEOPTERA, 203
groove, the surface, like the head, rufous and impunctate;
scutellum of the latter colour; elytra greenish-eeneous, with very
closely approached and irregular rows of rather fine punctures,
which are evenly distributed over the entire surface; below rufous,
the abdomen paler, the tibize and tarsi more or less black; the
prosternum rather broad, elongate.
Hab. Peru.
I know only a single specimen of this large species, which seems
allied to C. consularis Har. from Colombia, but differs in the
colour of the underside, the different comparative length of the
joints of the antenne, and the non-continued thoracic sulcus.
NASIGONA, gen. n.
Body elongate; antenne filiform; lower portion of the face
concave, the clypeus not separated; thorax transverse, the
anterior angles oblique, the surface obsoletely sulcate near the
base; elytra closely punctate-striate, their epipleure broad, con-
cave, the anterior and intermediate tibize unarmed, posterior tibize
with a small spine, non-sulcate, the metatarsus of the posterior
legs as long as the following joints together, claws appendiculate ;
prosternum narrowly elongate; the anterior coxal cavities closed.
The genus here proposed is allied to Oxygona, Nasidia Har., and
Systena, but differs from all of them in the structure of the head,
which at its lower portion resembles that of some genera of
Longicornia or of Loxoprosopus among the Halticide ; the structure
of the antenne and the shape of the thorax differ likewise from
Nasidia, and the punctate-striate elytra from Oxygona. From
Systena the genus may be separated by the structure of the
head and the very elongate joints of the antenne, as well as by
the punctate-striate elytra.
NASIGONA PALLIDA, sp. n. (Plate XX. fig. 12.)
Entirely pale testaceous ; antenne black, the apical three joints
testaceous, head and thorax impunctate ; elytra strongly punctate-
striate, the punctures nearly obsolete at the apex, a spot near the
scutellum and another near the apex piceous,
Var. Elytra without spots.
Length 5 millim.
Head entirely impunctate, testaceous, the frontal elevations
raised anteriorly and divided by an elongate fovea; clypeus forming
a single piece, perpendicular, with a feeble central and lateral
ridge, the anterior edge likewise narrowly raised ; labrum large,
apex of the mandibles black; antenne slender, all the joints,
with the exception of the second, very elongate, the basal one and
the last three joints testaceous, the others black; thorax twice as
broad as long, the sides distinctly constricted at the base, rounded
anteriorly, the anterior angles oblique, the surface impunctate ;
scutellum rather broad, impunctate; elytra wider at the base
than the thorax, the base slightly raised, strongly punctured in
closely approached rows, the interstices more or less distinctly
14*
204 MR. A. THOMSON’S REPORT ON THE INSECT-HoUSE. [Mar. 18,
longitudinally sulcate and slightly rugose or wrinkled, also more
or less distinctly punctured.
Hab. Chanchamayo, Peru.
Of the spotted form, a single specimen is before me ; these spots
are nearly round, one is placed at the base near the scutellum,
the other in a line near the apex. I look upon this specimen as
the normal form, although four others, contained in my collection,
have no elytral spots; there are no other differences between the
two forms.
EXPLANATION OF PLATE XX.
Fig. 1. Lactica costatipennis, p. 175.
» seminigra, p. 176.
5 bicolorata, p. 177.
. Disonycha angulato-fasciata, p. 188.
. Nephrica inelusa, p. 194.
b op brasiliensis, p. 196.
5 anifasciata, p. 197.
iS maculipennis, p. 192.
x claveri, p. 194.
10. s sanguinolenta, p. 193.
11. e staudingeri, p. 195.
12. Nasigona pallida, p. 203.
ODAD ME ws
March 18, 1902.
W. 'T. Buanrorp, Esq., LL.D., F.R.S., Vice-President,
in the Chair.
A series of mounted specimens of Insects reared in the Insect-
house during the past year was laid upon the table, and the
following report, drawn up by Mr. Arthur Thomson, the Assistant
Superintendent of the Society's Gardens, was read :—
Report on the Insect-house for 1901.
~The following is a list of the Lepidopterous Insects exhibited
i 1901 :-—
Silk-producing Bombyces and their Allies.
Asiatic.
Attacus atlas. Antherea mylitia.
cynthia. yama-Man.
ricint. Caligula japonica.
Rhodia fugax. simla.
American.
Samia cecropia. Telea polyphemus.
glovert. promethea.
ceanotht. Hypochera ‘0.
euryalus. Dirphia tarquinia
Attacus orizaba Eacles imperialis.
Actias luna,
_1902.] oN TRANSFORMATIONS OF SOUTH-AFRICAN LEPIDOPTERA. 205
African.
Nudaurelia cytherea. * Nudaurelia wahlbergi.
% zambesina. Gononuta postica.
tyrrhea.
Diurnal Lepidoptera.
Kuropean.
Papilio machaon. *Timenitis camilla,
podalirius. Charaxes jasius.
Thais polyxena. Vanessa antiopa.
American,
Papilio asterias. Papilio zolaicon.
ajac. Limenitis disippus,
cresphontes. ursula,
troilus.
Nocturnal Lepidoptera.
Acherontia atropos. Deilephila euphorbie.
Smerinthus ocellatus. Cherocampa alecto.
tilie. elpenor.
Sphinx ligustri. Ceratomia amyntor.
pinastri.
* New to Collection.
Of the Lepidopterous insects which I have the honour to
place before the meeting, the specimens of Vudaurelia zambesina,
NV. wahlbergi, and of a South African Cossus, presented by Mr. W.
_L. Sclater, are exhibited for the first time. The specimens of
Nudaurelia were received from near Pretoria. For the other
specimens of African Bombyces (which have been exhibited before
under the generic name of Antherw@a) the Society are indebted to
Messrs. W. L. Sclater, F.Z.S., H. W. Bell Marley, and Majors
Young & Clarke. For the cocoons of Dirphia tarquinia the
Society are indebted to Dr. A. E. Goeldi, C.M.Z.S., Para.
Mr. R. E. Holding exhibited and made remarks upon the skull
of a “Hummel,” or Hornless Stag, shot at Glen Tana, and the
skull of a Stag in which the left horn had been absent from birth ;
also the skull of a Welsh Sheep having four horns, a rare
occurrence in this breed; besides several specimens of horns of
Cattle showing bifurcation of the core-nodules in the horns and
other peculiarities.
Mr. R. Trimen, F.R.S., communicated a paper by Lieut.-Col.
J. M. Faweett, entitled ‘“‘ Notes on the Transformations of some
South-African Lepidoptera.” This memoir was in continuation
of one by the same author, already published in the Society’s
206 DR. H. GADOW ON THE EVOLUTION | Mar. 18,
‘Transactions’ (vol. xv. p. 291). It illustrated the earlier stages
of 32 species, of which 6 belonged to the Rhopalocera and 26 to
the Heterocera. As in the previous memoir, the Sphingide and
the several families of the Bombyces predominated in the series
illustrated, and many of these were of special interest in connection
with what was known of the earlier stages of the same groups of
allied species in the Oriental Region.
This Memoir will be printed entire in the Society’s ‘ Trans-
actions.’
The following papers were read :—
1. The Evolution of Horns and boners,
By Hans Gavow, M.A., Ph.D., F.R.S., F.Z.8.
[Received March 18, 1902. ]
(Text-figure 25.)
There are three works to which we naturally turn for infor-
mation concerning mammalian structures: Flower and Lydekker’s
‘Study of the Mammalia,’ Bronn’s ‘Thierreich, Mammalia by
Giebel, continued by Leche, and Gegenbaur’s ‘ Vergleichende
Anatomie der Wirbelthiere. But the treatment of the mor-
phology and phylogeny of the Ruminants’ horns and antlers in
all of them is singularly deficient and inadequate.
The actual development of Horns and Antlers has been studied
often enough, but no subsequent writer has taken the trouble
of sifting and reconciling the various contradictory statements.
Sandifort, in 1829, stated that the bone-core of the Bovine
horn is a compound structure, composed of a frontal outgrowth
or pedicle, and a superimposed ossification in a cartilaginous
matrix—the os cornu, which soon becomes indistinguishably
connected with the pedicle by synostosis, so much indeed that
the frontal sinuses in time extend not only into the pedicle, but
also into this os cornu. Lieberkiihn found cartilage in the
budding prickets of the Roebuck. Cartilaginous preformation,
with subsequent metaplastic ossification,was advocated also by
Joh. Miller, Gegenbaur, Kassowitz, and others. Landois de-
clared the development and ossification of the antlers as entirely
periosteal, Julius Wolff and Robin et Herrmann deny the
existence of cartilage, and call the ground-substance of the
budding antler amorphous embryonal tissue, or ‘substance pré-
osseuse.”
Riitimeyer, most careful observer and far-seeing thinker,
naturally homologized the os cornu of the Bovide with the
deciduous antler of the Cervidee; but the os cornu seems to have
fallen into oblivion until A. Brandt, as late as 1892, rediscovered,
or, rather, reinstated it. Brandt gives the following synoptic
table of “‘ Haut- und Knochen Hoerner” {@. ¢. epidermal and bony
1902. ] OF HORNS AND ANTLERS. . 207
“‘Horns”), an arrangement which we find adopted by Leche (op. cit.
p. 980) :—
Os cornu absent—Integument unaltered ............ the unpaired boss of the Giraffe.
Integument unaltered, os cornu persisting.,.paired horns of Giraffe.
Integument thin, short-haired, os cornu cadue?...... Antler sensu
Os cornu present strictiori—Subuli.
horn-sheath shed—Antilocapra.
es persisting —Bovide.
Integument cornified
Leaving aside the erroneous statement that the median “horn”
of the Giraffe possesses no separate bone-core or os cornu, but
only a frontal boss, this arrangement looks much better than
what it is intended to convey. In fact it represents exactly the
phyletic stages which I shall endeavour to prove as correct. But
Brandt says explicitly that the antler represents a higher, physio-
logically more complicated, differentiated stage of the bovine
armament. In this respect he agrees with Riitimeyer, who con-
sidered the Cervide as the “unfertigste und neueste” of all
Ruminants. Moreover, Brandt suggests the following evolutionary
stages: First, epidermal horns (analogous to those of Rhino-
ceros) ; secondly, the acquisition of frontal pedicles; thirdly, an
os cornu and an antler. Lastly, the same author doubts the
correctness of Riitimeyer’s reasonable view that the Antilocaprine
armaments are intermediate between those of Bovine and
Cervine.
Gegenbaur (1898), ignoring the os cornu altogether (op. cit.
p. 106-107), misses the proper connexion which exists between
horns and antlers. His remarks are not so precise and lucid as
they might be, and there is not a word about the most sug-
gestive preformation in a partly cartilaginous matrix.
Nitsche (1898) has tried to give a coherent account of the
whole question, but he does not seem to have made any other
than macroscopic examinations. Starting with well-ascertained
instances, observed in the Stag and Roe, where lesion of the side
of an otherwise normal pedicle has caused the growth of extra
antlers, he concludes that these cannot be anything but out-
growths or exostoses of the frontal bone. Consequently, he
argues, such extra antlers cannot be homologous with normal
antlers if such are dermal bones; 7. ¢., according to his unfor-
tunate definition, bones which are developed within the skin alone
and without connexion with the skeleton. He then refers to the
mode of growth of the first knobs and prickets of the young Roe,
that there is no difference in structure and development between
pedicle and pricket, anyhow that both are one and the same apo-
physis, while the separate existence of the os cornu in the Bovide
shows that their bone-core is an epiphysis. He then logically
opposes the two organs to each other. Moreover, he looks upon
the armaments of the Giraffe as exquisite types of cutis-ossification,
in fact, as examples of free dermal bones, since they develop
1 Obviously a misprint of Brandt and Leche for caducum,
208 DR. H. GADOW ON THE EVOLUTION [Mar. 18,
loosely in the skin, and even shift their position across the fronto-
parietal suture, and fuse very late with the cranial bones. He
naturally combats Riitimeyer’ s comparison of the antlers with
the Giraffe’s “horns,” which he seems to look upon as swi generas.
Not a few writers, amongst them Nitsche and Reerig, are not clear about the
meaning of the somewhat unfortunate terms “ Hautknochen,” dermal or membrane-
bones. Reerig, for instance, thinks that thereby are meant epidermal organs. In
reality they are contrasted with cartilage-bones as membrane-bones. ‘To call the
latter promiscuously dermal bones has caused endless confusion. A necessary
condition for ossification is the presence of an amorphous ground-substance or
matrix, which is then converted into, or rather supplanted by, bony tissue. Ossi-
fication is consequently always a secondary process. Unless the ground-substance is
preformed as amorphous embryonic tissue, it has first to be produced out of existing
adult cartilage or other connective tissue by the action of the osteoclasts or similar
katabolic, histioclastic cells, which by their breaking-down action upon the tissue
dissolve the latter into a medium in which osteoblasts can live, multiply, and by
excreting or attracting and arranging around themselves certain salts, turn into bone-
corpuscles. On the. surface of regenerating bone the marrow-cells, giant-cells,
myeloplaxes, seem to produce this eround- substance. In the case of cartilage this
is first destroyed, one might as well say dissolved, by the cells which immigrate
through the perichondrium, a process which happens frequently when membrane-
bone comes into contact with cartilage. It was a great step forwards when it
became understood that the place of origin of all bone- forming cells was to be
referred to the so-called basal membrane of the epidermis, whence osteoblasts
infiltrated or invaded the corium or mesodermal portion of the skin. Recent obser-
vations warrant us to go a step further, and to assume that the original home of all
skleroblasts was in the Malpighian layer of the epiderm itself. The oldest immi-
gration of skleroblasts from the ectoderm into cutis and other mesodermal tissue has
formed cartilage; the next immigration of skleroblasts has given rise to bone. The
latter being superior, supersedes the cartilaginous skeleton. ‘The ectoderm has by
no means lost the capacity of producmg either kind of skleroblasts. Extraordinary
excitement and requirements, reactions upon external stimulus, produce this rejuven-
escence, even in the mammalian skin.
Exquisite examples of true dermal bones are those ossifications “within the
skin” which in Amphibia and Reptiles are now generally called osteoderms. They
occur also, among Mammalia, in the Armadillos, but in no other group of this
class, unless it were in the Cetacea, where Kuekenthal has found traces of a dermal
armour. In the Amphibian Ceratophrys ornata the “dorsal shield,” although
very thick itself, has sunk in so deeply that it is now in contact with the vertebral
processes and is covered by the ordinary, movable skin. In Pelobates the skin of
the upper surface of the head is partly co-ossified with the underlying cranial bones,
giving them a pitted appearance. Now, frontal and parietal bemg membrane-bones,
or at least membranes which have received their bone from the cutis, this super-
imposed ossifying mass of Pelobates is a second instalment, or second generation of
dermal bone. Similar successive repetitions of the same process are demonstrated in
those Amphibian and Reptilian vomers which carry teeth, the vomers themselves
having resulted from the fusion of the basal portions of teeth which themselves are now
lost.—Concerning the cranial membrane-bones, there is no doubt that the original
cartilaginous roof has vanished (it is restricted to the dura mater), not because its
cartilage has been destroyed, or supplanted, by immigrating bony tissue, but because
it has been gradually suppressed by the approaching, investing, membrane-bone.
Similar instances of suppression, not conversion, are the greater portion of Meckel’s
cartilage, the premaxille and maxille, probably the palatine and quadrato-jugal
bones of Birds and Mammals, and to a great extent the mammalian quadrate through
its conversion into the os tympanicum. On the other hand, the human clavicle
surrounds, and is intermixed with, the precoracoidal cartilage.
The next points of importance considered by Nitsche are the
composition and shedding of the horny sheaths of Antilocapra,
which he does not homologize completely with the bovine horn-
sheaths. He prefers putting the Prongbuck’s horns into a
position intermediate between the velvet of the Giraffe and the
horn of the Bovide. The following is his terse summary :—The
1902. ] OF HORNS AND ANTLERS. 209
integument of the Giraffe is only hairy, that of the Prongbuck is
hairy and horny, that of the Bovide only horny. Having
satisfied himself about the apophysial nature of the antlers in
opposition to the epiphysial condition of the bony core of the
Giraffe and Bovide, he naturally feels justified about the com-
plete removal of the Giraffe from the Cervide and the funda-
mental difference between bovine and cervine ruminants. .
These conclusions are not satisfactory. rst, it is not likely
that several fundamentally different kinds of armaments should
have been developed within such a circumscribed and specialized
group as the higher Ruminants or Pecora. It is of greater
scientific value if we can trace all these armaments not only to
the same beginning, but bring them all into one evolutionary
line, so that these organs afford a clue to the phyletic develop-
ment of the various groups of Ruminants. The ultimate cause of
these armaments must have been the same, namely butting with
the head, causing irritation, which in turn leads to hypertrophy
of the cranial bones, together with the covering periost, cutis,
and epidermis’.
Secondly, the origin of the os cornu, sometimes with a separate
centre of ossification, has to be accounted for, since it is quite
impermissible to assume that it was a preformed bone in the
skin, which has later become fused onto the skull. Nor is there
a single instance In mammalian pathology of loose ossification in
the cutis due to repeated pressure or irritation, conditions which,
on the contrary and invariably, lead to exostosis of the underlying
already existing bone.
Thirdly, there is the paleontological evidence, not very plen-
tiful, but one fossil in hand is worth more as a proof of phyletic
speculation than ever so many ontogenetic observations. Curiously
enough, few paleontologists, whilst throwing much light upon
the evolution of the cervine antlers, have tried to take a broad
view of the phyletic side of the whole question, and the com-
parative anatomists have, as usual, left the fossils severely alone.
Lastly, there is the Giraffe and its relation, the Okapi, the
former with multiple, early appearing protuberances in both
sexes, the other apparently with none.
The question at issue requires a renewed investigation of the
development of the cervine and bovine armaments and a sifting
of the paleontological evidence.
I. Development of the Cervine Antlers.
The first growth of a buck’s antler in his first year is already a
compound structure. It begins with a bulged out elevation or knob
1 The general considerations and conditions have been well described and argued
out by J. J. Cunningham, ‘Sexual Dimorphism,’ pp. 73-104. The evolution of
horns and antlers, studied with due regard to the interesting physiological features,
is one of the clearest demonstrations of the acquisition and inheritance of new
organs, whilst any attempt to account for them (not their shape!) by variation and
selection alone would be ridiculous. :
210 DR. H. GADOW ON THE EVOLUTION [Mar. 18,
of the upper lamina of the frontal bone, which forms the pedicle.
This is a direct continuation of the frontal, identical with it in
its dense, lamellar structure, numerous Haversian canals and its
blood-supply, and it is covered by the same unaltered skin. It
is, in fact, an exostosis or apophysial growth. On the apex of
this pedicle the skin and the periost are thickened. The skin is
devoid of sudoriferous glands, produces no stiff, but only very
fine and soft, velvety hairs, is like them darkly pigmented and of
a glabrous appearance. The cutis is in direct, intimate continu-
ation with the periost, and contains numerous, but small vessels,
chiefly lymphatic, and only capillaries perforate the periost.
Immediately beneath it follows a dense layer of hyaline cartilage,
which, together with rapidly proliferating connective tissue,
makes up the apical portion of the pedicle and forms the growing
point of the future pricket. Vertical sections through the
growing pricket and pedicle show that the cartilage pervades the
top portion in the shape of strands, trabecule, and walls, which
partition off equally proliferating masses of ingrowing connective
tissue in which turn up bone-forming cells. The bulk of this
ingrowing tissue comes in with the vessels which extend from
the interior of the pedicle upwards into the base of the soft mass
on the top; little connective tissue enters together with the
small vessels of the periost. The process of ossification begins at
the base and near the periost, pervading the whole growth in the
shape of a very irregular framework, without forming concentric
bone-lamelle and with but few Haversian canals.
The first prickets or broaches are short-lived ; they are shed in
the middle, or even earlier, of the first winter. The shedding of
a full-grown antler has always rightly been referred to necrosis,
but it is a rather complicated process. To begin with, the antler
continues to ripen, or to harden, by the deposition of bone in the
more spongy, axial centre, long after the velvet has been frayed
off, the loss of which is consequently not the only, nor the main
cause of the decay of the antler. The latter is nourished not
only by the big vessels (branches of the temporal artery) which,
ascending in the skin and periost, cause the “ gutters,” but also
by the numerous vessels which ascend through the pedicle into
the interior of the antler. The base of the latter, where it passes
into the pedicle, becomes much denser and harder, instead of
remaining somewhat spongy in the core, and the blood-supply is
stopped. About the same time, ata level below this junction, 2. e.
within the top portion of the pedicle itself, the Haversian canals
are widened owing to activity of osteoclasts, and they become
confluent into a “vesorption-sinus.” This is met by a ring-
shaped furrow, which eats its way from the periost inwards. The
hardened base of the antler is slightly convex, while the resorp-
tion-sinus forms a somewhat deeper cup on the top of the pedicle.
Owing to this mode of resorption, which always affects the
pedicle, this becomes lower every year, but it makes up for this
loss by broadening. Long pedicles are consequently the older
1902.] OF HORNS AND ANTLERS. 211
stage, both onto- and phylogenetically. The cup is filled with
lymph, some oozing-out blood-clots and a rapidly increasing mass
of proliferating cells which are granulating from the walls of the
cup. This mass is soon, within a day or two, covered over by a
thin network of epidermal and connective tissue proceeding con-
centrically from the edges of the skin, which arises in the shape
of a thickened ring-wall. Large vessels, branches of the temporal
artery and facial vein, accompanied by branches of the facial and
trigeminal nerves, ascend in the much-thickened cutis which
covers the whole growth. These big vessels send only very fine
branches into the antler, and they soon become capillary. ‘These
seem to anastomose with the terminal capillaries of the vessels
which ascend within the antler. Owing to this arrangement, the
outer portions of the antler receive more calcareous salts than the
inner parts. They are denser, more opaque, and harder. Ossi-
fication of the whole soft and spongy mass proceeds from the base
and periphery upwards.
It is important to note that the preparatory process of shedding
follows immediately upon the time of greatest exhaustion, 7. e.
after the rutting-season, and that the beginning of the new
growth does not coincide with the awakening of sexual activity.
Herewith harmonizes the fact that adult stags, when castrated,
shed their antlers within a few weeks, whereupon a new growth
is formed, which, however, continues to grow throughout life,
resulting in abnormal, more or less monstrous antlers.
It is assumed generally that the fraying of the velvet has
originated through fighting, that the bared portion of the antler-
bone became necrotic, and had therefore to be renewed &c., and
that the whole process of stripping, necrotising, shedding, and
renewing has become rhythmical—a feature due to cumulative
inheritance. ‘This may be the case. But there is another con-
sideration. There would be no reason why antlers and velvet
should not grow continually, and mend or rebuild injured or lost
portions like other parts of the body, unless there occurs a
diversion or stopping of the energy and supply of building-up
material. Such a diversion is actually caused by the awakening
of the sexual glands. They are the important organs, and all
the energy and supply (which after all have their limit) not
necessary for the keeping up of the body and life of the animal
are concentrated upon the generative system, while nothing can
be spared for the further growth of secondary exuberances.
Therefore the blood-pressure in the head is diminished, the supply
of the skin covering the antler gradually ceases, and the velvet
itself becomes necrotic, from the apex downwards.
Il. Development of the Bovine “ Horns.”
The bovine or antelopine “ horn” is, as a rule, described as con-
sisting of a bony core, itself an outgrowth of the frontal bone, and
the horny sheath. In reality it corresponds exactly in growth
212 DR. H. GADOW ON THE EVOLUTION [Mar. 18,
and composition with the pedicle and antler of the Cervine.
The homologue of the antler has, since Sandifort, been called the
os cornu, and it forms by far the greater mass of the whole bony
cone. It has already been stated that it is continuous with the
pedicle, so much so that in many species of Cavicornia, especially
of Oxen, the sinuses of the diploé extend far into the cone, in old
animals far towards the apex. Occasionally the os cornu ossifies
from a special centre, separate from the frontal bone. It is,
however, short-lived as a separate entity. A. Brandt found it in
lambs as a small bony nodule, which could be lifted out of a corre-
sponding cup-shaped depression of the frontal bone in very young
animals. It fuses with the rest when the lamb’s horns are perhaps
3cm. long. Nitsche found it in the kid of a Chamois whose
horn-sheath was only 2 em. long, and he adds that it is already
firmly fused with the frontal bone in the first autumn. J myself
have several times come across specimens of ewe-skulls which had
been bleaching for a long time on the Welsh hills, and in which
the upper lamina of the frontal bone had fallen off at the precise
spot which normally carries the horns. There is no pedicle in
these specimens, but this absence is due to the feebly developed
state of the horns of these ewes, which as a rule do not carry such °
organs. In calves, even the youngest, a separate os cornu is
unknown, but they occur sometimes, pathologically, in polled or
hornless cattle as ever-growing bone loosely attached to the head
by the surrounding skin. It is a great mistake to imagine that
the occasional separate ossification of the os cornu is a primitive
feature.
The normal development of the calf’s head-gear is as follows :—
Slight elevation of the frontal bone into a comparatively broad-
based but low pedicle. This is surmounted by a shallow cone
composed of fibrous connective-tissue and cartilage. The cartilage
has its growing-point near the apex. The perisclerium is con-
tinuous with the periost of the pedicle portion. Ossification
proceeds from the pedicle upwards, transforming the soft growth
into a bony cone, the cartilage being gradually restricted to the
apex. The last traces of cartilage vanish when the horns are
between one and two inches in length, and the os cornu then
continues to grow by the ordinary subperiosteal mode, young con-
nective-tissue continuing to proliferate especially at the apex.
Blood-vessels enter from the pedicle and from the periosteum.
The periosteum passes imperceptibly into the rest of the cutis.
Vertical sections of a young “horn” half an inch in length
show all the minutest features in diagrammatic clearness. The
specimens were decalcified in picric and nitric acid, then cut and
stained with the triple stain of Erlich-Biondi, with hematoxylin
and picric acid, or with thyanine.
Outside is the dense mantle of horn passing towards the
Malpighian layer into the characteristic comb-shaped jagged
processes in the act of transformation into horn, each of the
processes resting upon a finger-shaped extension of a cutis-papilla.
1902.) OF HORNS AND ANTLERS, 213
Below and between them are numerous hairs, some of which reach
the surface and are imbedded in the horny mass, others are
hemmed in and suppressed.
The rest is best explained by a diagram. The important point
is that the cartilage, dense and hyaline, and in active proliferation
near the periost, changes further inwards into clusters of cells
which show the same features and the same thyanine-staining
action as the so-called muco-cartilage. Towards the strands of
young connective-tissue, which becomes more prominent as we
proceed inwards, the muco-cartilage cells show a breaking up of
their nuclei, so that only a glassy mass remains, interspersed with
débris of the cells and their nuclei. [When not decalcified, this
mass is somewhat opaque and bluish owing to infiltration with
calcareous salts. All this hyaline ground-substance is destroyed
by giant cells which are active on the margin and a little further
down. The place of the vanishing cartilage is taken up by the
network of connective-tissue strands, and in this appear very
active marrow-cells, and osteoblasts which build up trabeculze of
bone. Consequently the cartilage is not wanted at all for the
construction of the cone. This is also obvious from the fact,
mentioned above, that a few months after birth the cone continues
to grow long after the last trace of cartilage has vanished. The
cartilage is in fact an invader of dormant scleroblast-cells in the
periost, different from osteoblasts.] Concerning the horny sheath,
I have to mention two important points which have hitherto
escaped notice. First, the inclusion and gradual suppression of
hairs by the proliferating intercrinal horn-substance. Even in
old specimens of cattle-horns hairs become imbedded in the horn-
sheath, which in opposition to the bone-core always exhibits basal
growth. Secondly, the fact that calves, when several months old,
shed the first juvenile hornshoe. This is not always cast off in
one piece ; it may be frayed off, but this is a process very different
from the incessant wear and tear of the permanent horn-sheath
of the adult. The shed portion so to speak, the first generation
of the horn, is more porous, less solid than the permanent horn,
from the base of which it becomes separated to the extent of
several centimetres, as shown in the illustration (text-fig. 25,
p. 216). The whole process recalls the relation of the neossoptile
to the teleoptile or permanent feather, and still more the shedding
of our own feetal finger-nails.
II. Anéilocapra.
Not much more is known about the development of the Prong-
buck’s horns than what Forbes and, recently, Nitsche have
described. Nitsche has shown that the sheath is an aggregation
of sparse hairs connected by much “ intercrinal” horn-substance,
the whole process resembling somewhat the pathological ichthyosis
of calves. He has also shown that the prong is formed entirely
»
by the horn-sheath, and that there is no corresponding outgrowth
214 DR. H, GADOW ON THE EVOLUTION [Mar, 18,
on the bone-core. Forbes remarked that in his specimen “ the
prong is not yet visible, but may be felt at the base of the pedicel,
close to the skull, on the anterior margin of the horn.” (Cf. text-
fig. 25, III’.) In well-macerated specimens in the Cambridge
Museum, the difference between the rather long pedicle and the
decidedly short os cornu proper is well marked. Owing to the
continued and active growth of the shoe from the base, the axial
point and the prong are gradually pushed upwards, so that the
prong comes to lie far above the skull in a level even above the
base of the os cornu, which in the adult is thinned out into a
non-osseous, tapering, string-like cone of soft connective-tissue
and periost. The shoe continues to grow basally, and ultimately
engulfs not only the os cornu but also nearly the whole pedicle.
IV. The Giraffe and the Okapi.
It is of no importance to the present investigation whether the
few hitherto known skulls of the Okapi are those of young or
adult males or females. The skulls exhibit the same tendency
towards broad-based swellings on the fronto-parietal and facial
regions as in the Giraffe. Even in the latter genus these parts,
although slight and bulging, owing to the pneumatic condition of
the bones of such a weakly constructed skull, can ill be reconciled
with the only reasonable explanation of the genesis of horns and
antlers. We have to assume that the ancestors of the Giraffe had
stronger skulls, with serviceable antlers, and that these armaments
have caused the bosses of the supporting bones, and that in the
Giraffe these very armaments have degenerated into now merely
ornamental remnants, vanished in the Okapi. It is possible, as
Mr Thomas has sagaciously suggested, that the degeneration of
these armaments is correlated with the lengthening of the fore-
limbs and neck, the animals ceasing to fight with their heads and
using the powerful fore-limbs instead. This applies obviously to
the Giraffe, but not so easily to the Okapi, unless we look upon
the latter as the most degraded descendant of the whole group,
which, although perhaps never numerous, was certainly more
widely distributed in the shape of several genera and species.
At any rate, the Okapi represents not the beginning, but the most
modern and most modest member of a tribe which has flourished
in bygone times.
There are other proofs that the Giraffe’s armaments represent
no primitive condition. The bony growths appear loosely in the
skin, a condition which finds a parallel in the cases of separate
ossification of the os cornu of certain Bovine. Their matrix has
become so emancipated from the skull, that they shift their
position before fusing onto the cranium, and their mode of fusion
is most peculiar. As Mr. Thomas has expressed it graphically,
not only at the base of the growth, but around it, and quite
irregularly, there appear little bony nodules, which become
amalgamated with the cranium as if wax had been dropped upon
it, Such numerous, small and irregularly scattered “‘ osteoderms ”
1902. ] OF HORNS AND ANTLERS. 215
are not primitive, they are the expression of degeneration, of the
breaking up of a once powerfully developed bony growth.
Fossil Giraffe-skulls are unfortunately still unknown, but
Samothertwm with frontal and posterior protuberances, Sivatheriwm
with posterior growths hollow at the base and with frontal
growths, lastly Brahmatherium with posterior and frontal arma-
ments, are undoubtedly allied to the Giraffoid stock. We leave out
the unarmed Hydaspitherium and Helladotheriwm, which are
suspected by experts to be female Sivatheria. It has been ques-
tioned whether these armaments, huge, and sometimes branched,
were covered with skin and hair, or with horny sheaths. The
antlers of the original specimen of the male Sivatheriwm clearly
show deep and strong impressions of the blood-vessels, extending
almost to the tips exactly as in Cervine antlers. They were
undoubtedly covered with skin, and it would be hardly fair to
assume that this and similar specimens happened to have died in
the velvet stage. The same applies to Samotheriwm boissieri.
But it does not follow that these armaments were true antlers in
the sense that they were shed. Dr. Forsyth Major is inclined to
think that there are sutures between them and their base. If
there are any, they simply indicate the line of transition or
demarcation which is usually seen between the pedicle and the
rest. The hindmost pair of armaments of Sivatheriwm are hollow
at the base, a fact which speaks decidedly against periodical
shedding ; and the broadness of the base supports this view, since
a good and permanent vascular supply from below was thereby
ensured. The much shortened shape of the skulls of Sivatheriwm
and Brahmathervwm are unmistakable signs of specialization,
excluding the possibility that these huge creatures of the latest
Miocene or lowest Pliocene were the direct ancestors of Giraffes ;
but they were near relations and contemporaries.
We are now able to conclude that the evolution of Horns and
Antlers and similar cranial armaments has passed through the
following stages :—
I. Exostosis. Subperiosteal ostotic outgrowths of the cranial
bones, covered presumably with thickened skin-pads,
These armaments were multiple, occurring on various
parts of the skull. This type is rather old among the
Ungulata, witness the Kocene Amblypoda, e. g. Dinoceras.
It reoccurs amongst the Artiodactyla, which here alone
concern us. Protoceras of the Lower Miocene of Montana
is an almost ideal type in this respect, with its three or
four pairs of facial, orbital, and posterior bony excrescences
in the shape of uncouth ridges and neat cones. (Text-
fig. 25, I, p. 216)
II. Exostosis of the frontal bone producing a pedicle, with
epichondrosis of apical growth, which by subsequent basal
ossification becomes the antler. Skin originally unaltered,
hairy ; this and the chondrosteoma are shed periodically.—
Cervine type. (Text-fig. 25, II, p, 216)
216 DR. H. GADOW ON THE EVOLUTION [Mar. 18, .
These stages are repeated by every young Cervine male.
A portion of the integument is frayed off, at first perhaps acciden-
tally, then repeatedly during the annual rutting-time, and a
Text-fig. 25,
Hairs
SET; * Epiderm
¢ ¢ Cutis
——§£—S Frontal Bone
5; 41
Mh Cartilage V Cartilage & later on bone,ie Os Cornu
LY
———= Frontal bone
Evolution of Horns and Antlers.
I. Protoceras. Iil!. Antilocapra, adult; early stage.
II. Young Stag, with velvet. IIT. ye » 3 later stage,
Ila. Sivatherium. when the prong begins to
Il}. Giraffe. grow.
Ile, Okapi. IV. Domestic Calf.
rhythm of regeneration is established. The regeneration naturally
concerns chiefly the additional chondro-osseous apical portion, this
being the distal and therefore more easily injured part. But
1902. ] OF HORNS AND ANTLERS, 217
originally, as still shown by the mode of growth of the youngest
stage, no hard-and-fast line can be drawn between antler and
pedicle, and even now in recent species part of the bona jide
pedicle itself (7. ¢., that part which is not infiltrated with cartilage)
is annually destroyed and regenerated, although not “shed.”
It is safe to presume that the earliest Cervine had long pedicles
and short antlers, or rather prickets and breaches. The further
development into long and branched antlers is an instance of the
morphologically and pathologically well-known fact that organs
which are originally due to hypertrophic causes are liable to
grow toexcess. Thereis no maximum limit to the size of antlers
and to the number of tines in the Stag, although old individuals
are liable to ‘“ decline.”
The earliest typically Cervine creatures are referred to the
genus Paleomeryx. The somewhat mixed synonymy of genera
and species has to a great extent been unrayelled by Roerig, who
has described and figured every known specimen. Finality is,
however, impossible until we know for certain whether the
separately found pedicles and antlers, or both together, are suc-
cessive stages of one species, or represent the armaments of several
adult species, or genera, which did not pass keyond the respective
stages of broachers, forkers, Xe.
Two frontal pedicles and two pedicles with simple, low, spiked
antler-fragments are known from the Lower Miocene of Hessler.
They show already a slight burr, proof that the tips were shed.
Roerig, assuming that these fragments and the following speci-
mens form the successive stages or ‘‘ heads” of one and the same
species, refers them to Dicrocerus furcaius.
The second stage, or “ head,” is represented by typical broaches, -
with a distinct little burr, from the Dinotheriwm-sands. They
are referred hy Roerig to D. furcatus, second stage or head,
equivalent to D. elegans of Lartet = D. dicranocerus of Kaup.
The third stage, or head, with a thick, somewhat compressed
antler ending in a short fork, is D. furcatus from Steinheim, Mid-
Miocene. Another specimen, from the same locality, has a deeper
fork and a thicker burr—D. furcatus, fourth head. The buir,
not sharply marked off, but rather a thick swelling, bearsa striking
resemblance to a specimen of an immature Antilocapra in the
Cambridge Museum. The bony fork of course excludes any
further recemblance and affinity.
The last stage, with a gracefully forked long antler, with
typical burr upon the still long pedicle, is represented by D.
elegans from Sansan, Mid-Miocene; it is possibly the final head
of D. furcatus: synonymous with Procerrulus Gaudry, Miero-
meryx Lartet? The Neotropical Subulo s. Coassus e. g. C. rufus
still remains in the broacher stage; and Cervulus, the Muntjac, is
an incipient forker. Hydropotes inermis alone, of China, has no
outgrowth whatever.
The possession of deciduous, large, many-branched antlers
amounts to an enormous waste of energy and material during the
Proc. Zoou. Soc.—-1902, Vou. I. No. XV. 15
218 DR. H. GADOW ON THE EVOLUTION [Mar. 18,
life of the owner. Although full of grace and beauty, antlers are
morphologically very faulty structures, as wastefully contrived as
the shedding of the thousands of teeth of Sharks and Crocodiles.
The long duration of the growth of the antlers, their soft and
highly sensitive condition during this time, is even a distinct
trouble, not to say danger, a circumstance which shows clearly
that these organs are not primarily weapons to be used against
other species.
Ila. A side issue from II. Epichondrotic growths prepon-
derant, with multiple and broadened bases. Ossification
delayed but still proceeding from base. Cranial exostoses
or pedicles correspondingly reduced. These weapons with
an increasing tendency of intraperiosteal growth reached
a large size in width and length, and remained permanent
structures. The tips of the orbital and posterior pair of
weapons may have been covered with thickening epiderm,
more or less hairy; the bulk of the growth was perma-
nently covered with the unaltered hairy skin. It is
possible that this protecting cover and the tips of the
bony core were worn off without impairing the fighting
use of these massive structures, which need not die off
thanks to the remaining velvet, or (even if this was
ultimately lost) thanks to the unimpaired vascular supply
from the intertor of the broadened base. Creatures thus
armed reached their culmination in the huge Sivatherium
and Brahmatherium. (Yext-fig. 25, Ila, p. 216.)
Here we have to confess the existence of a painful gap and a
vagueness in connecting this type I] a with others of the main
line. This difficulty will remain until fossil ancestors of these
creatures are found. That they form a side issue is obvious enough.
So far as the few actually known genera are concerned, they
are of the latest Miocene, perhaps of the lowest Pliocene date,
anyhow considerably younger than bona fide Cervinee which we
ean trace back into Lower Miocene. In this respect, and by the
morphological agreement between the stalked posterior antlers of
Brahmatheriwm with pedicle and antler of an early Stag, we are
justified in looking upon this type Ila as a side-branch of the
main type II. On the other hand, the prevalence of multiple
outgrowth, facial, orbital, parietal, and generically variable, might
rightly be urged as a primitive feature, resembling m this respect
type I, so that the Sivatheriwm type would form a side issue
somewhere between I and JJ. However, it must be borne in
mind that multiple pairs of such weapons crop up pathologically
in recent Cervide, and normally even in the Antilopine 7etra-
ceras—facts which nobody can possibly consider as primary.
Consequently the multiple armaments of the Progiraffine crea-
tures are not absolutely an indication of their great phyletic age
and low position. (eters ; 3 ;
1902. ] OF HORNS AND ANTLERS. 219
IL6. Terminal, further development of type IIa. Epichon-
drotic growths proliferating freely and with broad bases,
so that they form intraperiosteal growths, separated from
the cranial bones, and consequently ossifying independently
of them, ultimately fusing with them. Cranial apophyses
or exostoses, or pedicles, much reduced in height. Disuse
of the outgrowths, implying cessation of the irritation
upon the basal periost (i.e. between the growth and the
cranial bone), explains diminution of the pedicles and
their late fusion and the long delayed process of ossification. °
But the development of the ecchondrotic mass, inherited
from the ancestral stock, and subsequent ossification still
go on, although without a purpose, and they produce
organs which, owing to their late fusion with the cranium,
their original home, now appear as osteoderms, although in
reality they are pseudo-primitive organs. The integument
remains hairy, except on the top where the epiderm
proliferates and cornifies a little. Example, the Giraffe.
(Text-fig. 25, IT 5.)
I1c. Apparent loss of all these armaments, the last remnants
being frontal bosses: Okapi. (Text-fig. 25, IIc, p. 216.)
Tt is worth noting that, while the females of Stvatherium and
Samotherium are, by general consent, not credited with “antlers,”
the Giraffe makes an exception in this respect. This fits in with
the view, expressed in this paper, that Giraffes represent a
terminus of one line of development. There are some typical
Cervine of which both sexes are antlered. The acquisition of
secondary sexual organs by the females is mainly a question
of time. It is an illustration of simple, direct inheritance from
the other sex, so common in organs which are connected with
sexual activity, e.g. clitoris, mamme, spurs. These things are of
not the slightest good to their new possessors, but they do no
harm either. They are therefore neglected, rather not discovered,
by natural selection.
III. The same initial stage as type IT. A long pedicle with a
simple broach, covered with hairy skin, but the epidermal
portion of this tegumentary sheath proliferates, glues the
hairs together and embeds them. The horny sheath
is an efficient protection against injury; the external or
cutaneous and the internal vascular supply remain, and
the simple antler is shed no longer. Immature specimens
still show a thickened, burr-like swelling at the juncture
of the pedicle and antler. We assume that the horn-sheath
consisted originally of an imperfectly welded material still
‘liable to fraying, until 1t became effective enough to pre-
vent any necrosis and subsequent shedding cf the antler,
which thereby becomes an os cornu. So long as the hair
preponderates in the deeper strata, the ue and
Lo
220 DR. H. GADOW ON THE EVOLUTION [ Mar. 18,
renewal of the hairy coat is likewise repeated by the horny
sheath. This stage is still represented by Antilocapra,
although the horny sheath by continued basal growth
gradually envelops also the greater part of the pedicle.
(Text-fig. 25, III, p. 216.)
ITV. Direct continuation of types II and III, still repeated
stage by stage ontogenetically. An improvement towards
the preponderance of the intercrinal horn-substance, the
conversion of the sheath into a morphologically well-
finished horn-sheath, the suppression of the hairs, and of
periodical shedding of any part of the whole compound
weapon, was only a question of time with onward evolution,
This, the highest and most perfect stage, is represented by
the typical Antelopine or Bovine Ruminants, of which
their peculiar member, the Prongbuck, still falls short.
They are morphologically the highest, paleontologically
the latest of Ruminants. Herewith it agrees that horns
are carried by both sexes, whilst the inheritance of these
organs by the females is still a rare exception amongst the
Cervine. Moreover, these weapons, having become per-
‘manent and evergrowing, and therefore useful throughout
the year, are of much greater value to their bearers.
(Text-fig. 25, IV, p. 216.)
Attention has already been drawn in this paper to the important
fact that the horns of a young calf still contain a considerable
number of hairs mixed up in the sheath, and that in older
animals such hairs are restricted to the more basal portions;
secondly, that the top cone of the hornshoe is shed. In Ewes
this first generation falls off as a thin, transparent cap of the
size and shape of half a hazel-nut. In fact this first cap of the
Bovine horn is in every respect homologous with the shedding
sheath of the Prongbuck. The Oxen, Sheep, and Goats now
exhibit only once a process of shedding which in their immediate
ancestors must have been of frequent occurrence, and which in
the Prongbuck is still a periodical feature.
The types I, II, III, and IV, exemplified by the Eocene
Dinoceras, the Cervine since the Lower Miocene, the Prongbuck,
still existing, and the hollow-horned Ruminants or Bovine, are
an illustration of onward phyletic evolution; and these stages
are still faithfully repeated in the development of the recent
species. Ontogeny is a shortened recapitulation of phylogeny.
Titles of the more important Literature referred to in the text.
Sanpirort, G. Over de Vorming en Ontwikkeling der Horens
van zogende dieren in het algemeen en van die der Herten-
beesten in het bijzonder. Nieuwe Verhandl. i. Klasse
Koningl. Nederl. Inst. Wetenschap. ii. (1829). pp. 67-106 ;
with 7 plates. «
1902. ] OF HORNS AND ANTLERS. - 221
GncgenBAur, CO. Ueber die Bildung des Knochengewebes, Jena.
Zeitschr. 1867, p. 206.
Vergleichende Anatomie der Wirbelthiecre, i. (1 898), p. 107.
Lieserkiipn, N. Ueber den Abfall der Geweihe und seine
Aehnlichkeit mit dem carioesen Process. Arch. f, Anat. u.
Phys. 1861, pp. 748-759, pls. 18, 19.
——. Ueber Wachsthum des Stirnzapfens der Geweihe. Arch.
f. Anat. u. Phys. 1865, p. 404. (More elaborate in Sitzker.
Ges. naturf. Freunde, Berlin, 1865, p. 9.)
Lanpots, L. Ueber Ossification der Geweihe. Centralbl. medizin.
Wiss. 1865, No. 16, pp. 241-243. :
Jospru, C. A. Gehoernbildung des . Rehbocks. Monatschr.
Forst- und Jagdwesen, xix. (1875), pp. 304-313.
Ritimsyer. Beitrige zu einer natiirlichen Geschichte der
Hirsche. Abhandl. Schweizer. Paleont. Ges. vii. (1880),
x. (1883).
Forses, W. A. Remarks on the Horns of the Prongbuck. P. Z.8.
1880, pp. 540-543.
Rosin et Herrmann. Mémoire sur la Génération et la Reé-
génération de l’os des Cornes caduques et persistentes des
Ruminants. Journ. d’Anat. et de Physiol. 1882, pp. 205—
265, pl. xix. (Excellent account of the development in the
Roebuck.)
Branpt, A. Ueber Hoerner und Geweihe. Festschrift f.
Leuckart, 1892, pp. 407-413.
Nirscuse, H. Studien tiber Hirsche. Pt. 1, Leipzig, 1898.
Rorric, A. Ueber Geweihentwickelung und Geweihbildung.
Arch. f. Emntwickl. Mechanik, x. (1900), pp. 525-644,
pls. xii-xili.; xi. (1901), pp. 65-148, pp. 225-309,
| Numerous drawings of antlers, both normal and abnormal ;
with a complete list of the literature and a historical review
of the whole subject, but he, as well as Brandt and Nitsche,
have restricted themselves practically to macroscopic fea-
tures. |
Important from a general point of view are also :-—
Sir Victor Brooke. On the Classification of the Cervide...
P. Z.S. 1878, pp. 883-928.
Scutosser. Beitrag zur Kenntniss der Stammesgeschichte der
Hufthiere. Morph. Jahrb. xii. p. 1 (1887).
Cunnincuam, J.T. Sexual Dimorphism in the Animal Kingdom,
London, 1900.
P.S.—On the day this paper was read Dr. Forsyth Major
gave me intelligence of a paper by Dr. J. Ulrich Diirst. A copy
of his “ Versuch einer Entwicklungsgeschichte der Hoerner
der Cavicornia nach Untersuchungen am Hausrinde” | Forsch-
ungen auf dem Gebiete der Landwirthschaft, Frauenfeld, 1902]
reached me on March 24th. The author has also observed the
shedding of the first horn-sheath; he likewise correctly states
222 MR. R. I, POCOCK ON A NEW | Mar. 18,
that the budding growth of the bone-core differs in structure and
mode of development from the frontal bone proper, but he
emphatically doubts the temporary separate existence of the
os cornu, and he feels satisfied that it is not formed by the interven-
tion of cartilage, since the substance in question was not coloured
blue by hematoxylin staining. He and others will have to
accustom themselves to the existence of cartilage in places where
text-books carefully abstain from mentioning it.
2, On a new Stridulating-Organ in a Scorpion.
By R. I. Pococx, F.Z.8.
[Received February 25, 1902. ]
(Text-figure 26.)
Stridulating-organs have been found in three genera of Scor-
pions, viz., the large species of the Oriental Region and Tropical
Africa referred to Palamneus and Pandinus, and the South-
African members of an allied form Opisthophthalmus’. In the two
first-named the organ lies between the basal segments of the
chelz and of the legs of the first pair; in the latter between the
inner surfaces of the mandibles or their upper edge and the front
border of the carapace. In all three cases it consists in the main
of peculiarly modified bristles. No organ of similar function has
as yet been discovered in any other family of Scorpions. But in
the Buthoid genus known as Parabuthus, which ranges from the
shores of the Red Sea to Cape Colony, I find a stridulator (text-
fiz. 26, A & B, p. 223) differing entirely both im structure and
position from that of the Scorpions above mentioned.
It has long been known that the upper sides of the proximal
segments of the tail in Parabuthus are furnished in the middle
with an aggregation of granules, so fine and close-set as to be
appropriately comparable to shagreen, The granules are some-
times thickest and coarsest in the median groove, but finer and
more scattered at the periphery of the area; sometimes of
uniform strength throughout: sometimes they are confined to
the median groove; sometimes, and more often, they encroach
upon the adjacent area of the surface that bears them.
Of the species known to me, the granulation reaches its highest
point of development in Parabuthus flavidus Poc., where the
granules have run together across the middle line to form short
parallel transverse ridges with their free edges directed backwards
(text-fig. 26 B).
The surface that bears this granulation also differs in forma-
tion according to the species. In the more northern and less
specialized forms—such as P. liosoma, P. abyssinicus, P. hunteri,
P. granimanus, and P. heterurus—the area in question is but
little modified, remaining normally depressed and grooved in the
1 Pocock, Nat. Science, ix. pp. 17-25 (1896), and Ann. Mag, Nat. Hist. (6) xviii.
pp. 75-77 (1896),
1902.] STRIDULATING-ORGAN IN A SCORPION, 223
middle line; but in many of the southern types—e. g., P. plani-
manus, P. neglectus, P. villosus—the whole upper surface of the
segments tends to become flattened and horizontal both in a
longitudinal and transverse direction. A similar granular field
is developed between the dorsal keels on the last abdominal
tergite.
Text-fig. 26. B
.
SS
Sw
=
SOs
Stridulating-organ of Parabuthus flavidus.
A Lateral view of tail, to show the action of the sting during stridulation.
B. Dorsal view of last somite of abdomen and of 1st and 2nd caudal segments,
showing the ridges on the median groove of the two segments.
If the tail of one of these Scorpions be brought into the attitude
usually assumed by these animals when striking, and the point of
the sting be scraped over the granular field, a very distinct sound
is emitted, resembling that produced by drawing the point of a
needle over fine sand-paper ’.
There is as yet no direct evidence, based on observation of the
living animal, to prove beyond dispute that these granules have
the function here assigned to them, but the facts which support
the conclusion are the following :— we
(1) The sound can be artificially produced, and is audible to
me at a distance of ten yards or more.
(2) The scorpion itself is capable of performing all the move-
ments necessary for its production,
1 A similar but less complete development of granules, subserving no doubt the
same purpose, is observable in two North-African species of Buthus—B. bicolor
and B. eneas.
224 - DR. R. BROOM ON THE ORGAN OF [ Mar. 18,
(2) The granules are especially well-developed upon the first
and second caudal segments and upon the last abdominal tergite,
against all of which the point of the sting can be forcibly scraped.
On the third caudal segment, upon which the sting is capable of
but little movement, they are scarcely or not at all developed, and
upon the fourth and fifth, which cannot be touched by the point
of the sting, they are absent.
(4) The longitudinal flatness of the granular area on the first
and second caudal segments, which results from the uprising of
the groove and the elevation of the anterior part of the upper.
surface, can be explained on the supposition that rt is designed to
give the sting a long and continuous sweep from segment to
segment, without the danger of catching against their posterior
edges or of wounding the arthrodial membrane. It is difficult to
see what other interpretation is to be put upon this special and
unique modification of the segments in question.
3. On the Organ of Jacobson in the Elephant-Shrew
(Macroscelides proboscideus). By R. Broom, M.D., B.Se.?
[Received February 4, 1902.]
(Plate XX.)
From the examination of the organ® of Jacobson in a large
series of mammals, I, in 1897, concluded that it varies surprisingly
little in even very dissimilar genera of a common Order. In the
Marsupialia the chief Polyprotodont genera have their organ of
Jacobson very much alike, while even in the Diprotodonts the
organs are all formed on a type which differs but little from that
found in the Polyprotodonts. While in all the Rodents, so far
as examined, the organ is formed on a single peculiar type which
seems to be a modification of that found in the Marsupials, in the
higher mammals a single type of organ is found in forms so varied
as the Hedgehog, Bat, Lemur, Cat, Sheep, and Pig.
Tt would thus appear that the organ of Jacobson is but little
influenced by the habits of the animal, that it remains a clear
indicator of the early family relationships of a genus when
almost all the other ancestral characters have been so modified
as to be scarcely recognizable, and that hence it is of consider-
able importance in determining the precise affinities of aberrant
mammals.
Having recently had oceasion to make a series of sections of
the snout of a fetal Elephant-Shrew (acroscelides proboscideus),
in connection with a study of the development of the palatine
process of the premaxilla, I was naturally much interested in
observing the condition of Jacobson’s organ, especially as W. K.
! Communicated by Prof. G. B. Howes, F.R.S.
2 Hor explanation of the Plate, see p. 227.
3 'Traus. R. Soc. Edinb. vol. xxxix. p. 234,
FP. Z.S. 1902, vol L Fl XXII.
\ Ce
DP)
Parker & Wed inp.
ME. Parker lith.
EB. del.
>
JACOBSONS ORGAN IN MACROSCELIDES.
i) ‘
eae
ie Wy, em vl
aailees ny
en
i ie i
ren Uae Lf
1902. ] JACOBSON IN THE ELEPHANT-SHREW. 225
Parker has shown that in the allied genera Petrodromus and
Rhynchocyon there are a considerable number of Marsupial
characters. In the Hedgehog the organ is formed on the common
Eutherian type, and I expected to find in Macroscelides indications
of marsupial affinity. When the organ was investigated, however,
it was seen to be quite different from that in any Eutherian
hitherto examined, and to be typically Marsup‘al in almost every
respect.
Before entermg upon comparative observations, it will be con-
venient first to describe the condition of parts in Macroscelides.
Tf a section be made near the middle of the proboscis, it will be
seen (Pl. XXI. fig. 2) to be formed of a ring of cartilage (a.n.),
enclosing the two nasal passages and surrounded by soft tissues—
muscles, tendons, and skin. The skeletal portion is made up of a
well-developed median nasal septum (7.s.) and two alinasals (@.n.),
which sweep round from the upper end of the septum and meet
each other inferiorly. From this lower point of union of the
alinasals they pass upwards to meet the lower end of the septum.
From the inner side of each alinasal, near the level of the base of
the septum, there passes inwards a small turbinal which is an
anterior continuation of the inferior nasal turbinal; and from the
point where the incurved end of each alinasal meets the base of
the septum there passes downwards and outwards a second
turbinal plate which may be referred to as the septal turbinal (s.t.).
With very little modification, this description might refer to
any section of the proboscis. On approaching the anterior end
(Pi. XXI. fig. 1), however, the turbinals are found to be less
developed, the nasal septum slender and fused with the alinasals
inferiorly, and the upper half of the alinasals to be separated
from the lower. Near the anterior na:al opening a transverse
section shows a pair of alinasals above, a pair of cartilages
on the nasal floor, and a pair of rather complicated lateral
cartilages which apparently form nasal valves. The anterior
nasal opening looks outwards and slightly downwards.
In a transverse section near the root of the proboscis, the only
noteworthy differences from the more anterior sections are that
the base of the septum is considerably larger, while the lower
halves of the alinasals are not only separate from each other,
but are distinct from the alinasals above—forming nasal-floor
cartilages.
When the plane of the premaxilla is reached (Pl. XXI. fig. 3),
the outer and lower part of the nasal-floor cartilage becomes lost,
_ only the part situated immediately below the base of the septum
and which forms the septal turbinal remaining.
A few sections in front of the plane where the palatine process
is given off from the premaxilla, the premaxilla ( pmax.) sends
upwards a narrow plate as a support to the inner side of each of
the two cartilages which lie at the base of the septum. These
plates form the anterior ends of the palatine processes (p.p.).
About this same piane the smaii cartilaginous plates, which in the
226 " DR. R. BROOM ON THE ORGAN OF [ Mar. 18,
more anterior sections have been seen to form the septal turbinals,
become detached from the inner plates.
A section immediately behind the anterior end of the palatine
process shows the inner plate of the nasal-floor cartilage dipping
down in the cleft between the palatine process and the premaxilla,
supported on its inner side by the vertical plate of the former.
The papilla is fairly large, and the naso-palatine canal is seen
passing upwards and inwards by its side.
A few sections beyond this plane we see (Pl. X XT. figs. 5 & 6)
that the na:o-palatine canal (z.p.c.) on passing further upwards
turns outwards as it opens into the nasal cavity. On its upward
passage it receives the duct of Jacobson’s organ (/.0.). This duct,
which lies almost vertically, passes between the large vertical plate
and the apparently detached outer portion of the nasal-floor
cartilage and opens into Jacobson’s organ near its anterior end.
The organ extends very slightly in front of the point where the
duct is given off.
Immediately behind the duct (Pl. X XI. fig. 7), the lower end
of the vertical plate becomes attached to the outer, apparently
detached, portion, forming a floor to the organ. It will thus be
seen that Jacobson’s cartilage has an outer bar exactly as in
Marsupials.
The Jacobson’s organ itself is of moderate length and presents
no features of special interest in the foetus. The posterior part
of it appears to be devoid of sensory epithelium and to be merely
the duct conveying the secretion from a large number of glands.
In the adult, the cartilages are essentially similar in arrange-
ment to those in the fcetus, but an additional feature is to be
observed in the presence of a well-developed cartilage in the
papilla (Pl. X XT. fig. 8). The organ is in section (Pl. XXI.
fig. 9) somewhat kidney-shaped, with a single large vessel running
along the hilus. The sensory epithelium is confined to the inner
wall, and the organ is abundantly supplied with glands (g.).
From the above description it will be seen that in its relations
the organ has little or no resemblance to the highly specialized
type met with in most Eutherians, and that all its peculiarities
are those typical of Marsupials.
In the Marsupial the following may be regarded as the most
typical features of this region of the head:—(1) The Jacobson’s
organ opens into the naso-palatine canal near the poimt where
the canal opens into the nasal cavity; (2) the anterior part of
Jacobson’s organ is protected externally by a cartilaginous bar
which passes from the outer edge of the lower part of Jacobson’s -
cartilage behind to the outer edge of the upper part in front;
(3) the naso-palatine canal is never supported by a cartilaginous
process from the nasal-floor cartilage; (4) the nasal floor has no
cartilaginous support behind the region of the naso-palatine canal ;
(5) the papilla has a well-developed cartilage; and (6) the Jacob-
son’s organ has usually a single large vessel running along its
outer face.
1902.] JACOBSON IN THE ELEPHANT-SHREW. 227
While almost all Marsupials exhibit these features, no higher
mammal hitherto examined agrees with the Marsupials in more
than three of these characters. The peculiar condition of Jacob-
son’s cartilage, whereby a cartilaginous bar is present along the
outer wall of the anterior part of the organ, and which I regard
as a remnant of the turbinal of Jacobson’s organ retained in the
Monotremes, is among the Eutheria only met with in the Eden-
tata (Dasypus), and in a rudimentary condition in some Rodents.
Only the Hdentates again agree with the Marsupials in the
absence of a cartilaginous support to the naso-palatine canal.
The presence of a cartilage in the papilla, though occurring in
most Marsupials, has hitherto only been observed among higher
mammals in the little Bat Iiniopterus, and possibly as a rudiment
in Cavia.
From the fact that Maeroscelides agrees with the Marsupials in
every detail of the anatomy of this region, we are forced to the
conclusion that it is a very near relative of the Marsupials, and
has probably very little affinity with the more typical Insectivores.
That the marsupial characters are not confined to the nose we
know from Parker’s work. In the tympanic region and in the
remarkable condition of the orbito-sphenoid the marsupial affinities
are quite as remarkable.
It is further interesting that, while Wacroscelides in the anatomy
of its anterior nasal region agrees more closely with Perameles
than with other Marsupials, Perameles in one or two respects
agrees more closely with J/acroscelides than with most of the
typical Marsupials. The striking similarity of structure is shown
in the drawing (Plate XXI.), where a section of the anterior part
of Jacobson’s organ in Perameles is shown (Pl. XXI. fig. 10) for
comparison with the section of J/acroscelides (fig. 8).
It may be noted that IMacroscelides has a discoidal deciduate
placenta, and that the young are born in a well-developed con-
dition.
EXPLANATION OF PLATE XXI.
References.—a.J.c., anterior portion of Jacobson’s cartilage; a@.J.o., anterior por-
tion of Jacobson’s organ; a.., alinasal; a.p.p., anterior spur from palatine process ;
g., glands; i.c., smail isolated cartilage—possibly a rudimentary Stenson’s cartilage
(not indicated in the adult) ; 7.¢., inferior turbinal; J.c., Jacobson’s cartilage; J.0.,
Jacobson’s organ; Ud., lachrymal duct; w.g., nasal gland; m.v., maxilla; x.p.c.,
naso-palatine canal; z.s., nasal septum; 0.6.J.c., outer bar of Jacobson’s cartilarge ;
p.c., cartilage of papilla; pma., premaxilla; p.p., its palatine process; s.¢., septal
turbinal,
Figs. 1, 2, 3, 4. Transverse section of snout of fetal Macroscelides proboscideus.
X 24.
Figs. 5, 6, 7. Transverse section of the same region of naso-palatine canal. xX 40.
Figs. 8,9. Transverse section of Jacobson’s organ in adult Macroscelides. X 18.
Fig. 10. Transverse section of anterior end of Jacobson’s organ in Perameles nasuta
(young). xX 17.
Addendum (March 1902).—While writing this paper I had
taken for granted that the cranial characters described by Parker
in Rhynchocyon and Petrodromus would be common to the other
228 MR. F. CHAPMAN ON FORAMINIFERA AND [ Mar. 18,
genus of the family, more especially as the inter-relationship of |
the three genera is apparently very close; but, on looking into
the structure of the skull, I find that in both Macroscelides pro-
boscideus and M. rupestris there is a distinct optic foramen. The
marsupial characters of the tympanic region are, however, as
marked in Macroscelides as in the other genera.
4. On some Foraminifera and Ostracoda from Cocos Keeling
Atoll, collected by Dr. C. W. Andrews, 1898. By
FrEepERICK CHAPMAN, A.L.S., F.R.M.S.
[Received February 25, 1902.]
(Text-figures 27 & 28.)
On his return from Christmas Island in 1899, Dr. Andrews was
good enough to submit to the writer some sands gathered between
tide-marks in the Cocos Keeling Islands for examination. A
casual glance at the material was sufficient to prove it worth
recording, and more especially since the samples were taken both
from the lagoon and from the outer side of the atoll.
Though far removed geographically, the microzoic fauna of
Funafuti bears some striking analogies with the present collection,
chiefly on account of the similarity of conditions in the habitats
of the two faunas.
An especially noteworthy feature with regard to the gatherings
now described is the frequent occurrence of the rare and
occasional form Pavonina, chiefly on the lagoon side.
With respect to the source of these samples, Dr. Andrews
informs me that those from the outer reef came from a spot at
some distance from the transverse channels which communicate
with the lagoon. The lagoon material came from the inner
margin of the reef, and it would most likely be commingled to
some extent with organisms washed in through the sea-channels ;
but the general facies of this series, however, points to their
having lived in sheltered water.
The species of Foraminifera are numerous for such a small
gathering, amounting in all to 76. Some short notes are added
regarding those which are of especial interest, either on account
of their rarity elsewhere or their exceptional development. None
of the forms appear to be actually new, but there are many
peculiar modifications in form.
The Ostracoda number 28 species, and include two new forms.
They are nearly all well-known littoral species, and are fairly
equally distributed both inside and outside the lagoon.
Notes on the Ostracoda from Cocos Island.
The genus Badia is represented by 8 species, one or two of
* Communicated by C. Davies SHERBORN.
1902. ] OSTRACODA FROM COCOS KEELING ATOLL, 229
which have occurred in abyssal deposits, such as B. milne-edwardsir
and B. crosskeiana. They are all, however, of far more frequent
occurrence in shallow-water dredgings. A few of the species
have a wide geographical range, as 8. milne-edwardsi, b. acanthi-
gera, and B. crosskeiana, which are also northern species. Of the
10 species of Cythere two are known from northern areas, namely,
C. prava and C. stimpsoni. Anotber species, C’. dictyon, is more
often found in deep water, being recorded from fifteen out of
twenty-five of the ‘Challenger’ dredgings at depths of 1000
fathoms and more. The remainder are well known from shallow-
water dredgings. The genus Loxoconcha is represented by four
species, three of which here a fairly wide range, namely, LZ. alata,
L. honoluliensis, and L. anomala. There are three species of the
genus YXestoleberis, two of which have a wide range; one of them,
X. depressa, is also common in dredgings off the British coast,
from the North Atlantic, and also as a post-Tertiary fossil from
Scotland, Ireland, Norway, and Canada. The only species of
Cytheropteron recorded here, namely C. longicaudatum, was origi-
nally described by Dr. G. 8. Brady from material dredged in the
Fiji group.
Description of New Species of Ostracoda.
(The specimens were not preserved sufficiently well to enable
the organisms to be examined.)
CYTHERIDEIS ANDREWSI, sp. nov. (Text-fig. 27.)
Carapace suboval, elongate, somewhat arcuate and compressed.
Dorsal margin, in side view, rather irregularly curved; ventral
margin concave and sinuous. Anterior extremity produced ; :
Text-fig. 27.
Cytherideis andrewsi.
a, right valve, lateral view ; 6, edge view; ¢, end view. X 42 linear.
posterior evenly and broadly rounded. Edge view compressed
ovate. End view subcircular. Surface of the carapace covered
with fine pittings; and the muscle-spots in the median area
disposed in rosette-form. Length -57 mm.
Cocos Island, lagoon; very rare.
This species differs from C. levata G. 8. Brady in form and
also in the surface-markings on the carapace itself; but it is
evidently allied in some respacts,
230 MR, F. CHAPMAN ON FORAMINIFERA AND [ Mar. 18,
CYTHERELLA VESICULOSA, sp. nov. (Text-fig. 28.)
Carapace subrectangular, elongate, seen from the side. Surface
of carapace highest near the dorsal margin, sloping towards the
ventral, with a depressed area in the middle of the valve near the
ventral margin, in the lowest part of which there is a circular pit.
Text-fig. 28.
Cytherella vesiculosa.
a, right valve, lateral view; 5, edge view; ¢, end view. X 42 linear.
Anterior part of carapace gently sloping towards the margin;
posterior border steep. Edge view cuneate. End view oval.
Surface of valves ornamented with numerous deeply-set pittings
or cavities. Length -76 mm.
Cocos Island, lagoon; rare.
The nearest form to this handsome species seems to be C. semi-
talis G. S. Brady; but it differs in having only a part of the
carapace covered with pittings, which are not, however, exactly
comparable with those in our form, and the carapace itself is
also shorter and stouter. The ventero-median pit is also wanting
in C. semitalis.
Ostracoda from Cocos Island.
Species. Seaward face. Lagoon.
1. | Bairdia tenera G.S. Brady ............... «. Y. Yr.
Px 5 ventricosa G.S. B. ......... eee TOME Met) ig eee?
3. “A P acanthigera G.S. B._.........5-- Moai Yr.
4. 3 milne-edwardsii G.S. B. ......-+- Pee hal) gp eee
5. 5 CHAHADUIGHA® (Cts 1B congnpencesosce col - cdodoc v.r.
6. F5 amygdaloides G.S. Be ......-00- | eee C.
ie crosskeiana G.S. B. ...... eee f. V.C.
8. a woodwardiana G.S. Be .........c | tees 1%
9. | Cythere scintillulata G.S.B. «2.0.0... |e at OVE A Sb
10. $5 cancellata G.S. Bw ...... ccc eee eee VeTates eye Wyre teemesste
11. 3 SHR Crsts\s 18s gancoodoansanecbes suc V.1. v.r.
12. 5) cristatella G.S. Bu. oo... cece eee Tea ella F eiemuareeres
13. én rastromarginata G.S.B. ......f 6 sees v.r.
14. 5 auaraniiana GSE acococccnccrauscoa|| 1» dccooc v.r.
15. Bs wyville-thomsoni G.S.B. ...ccec| eevee v.r.
16. re GAUL OOO (Cigisb | B}baqnnopessondeosoosecd||) | coabdc Yr.
f., frequent ; ¢., common; v.e., very common; Y., rare; v.r., very rare.
1902. ] OSTRACODA FROM COCOS KEELING ATCLL. 231
Ostracoda from Cocos Island (continued).
Species. Seaward face. Lagoon
17. | Cythere prava Baird . v.I. v.r.
18.- * dictyon G.S. B. Sect neReR esis f. Gp
19. | Loxoconcha alata G.S. Bw... ccc cee ee f. ®,
20. a GHOMALGA GAS bee eee VAT eller Se eee
21. 3 honoluliensis G.S. B. Mets ce ieee vets
22. 5 avellana G.S. B. Bie Ved tights Tee MARL
23. | Xestoleberis depressa G.O. Sars Ch ae eee
24, = GUTAGD Cia tSo 1B, scon0sas0o0960900 Warsed 2'P Rey Bee
25. TOCRRUMMOGEE (Cats Wkeassnesccassl] dvd vr
26. | Oytheropteron austin © Gass bene v.r. vr.
27. | Cytherideis andrewst, sp. nov. Sa5éal NU. Von vr.
28. | Cytherella vesiculosa, sp. nov. Bapecenea | eh eeesa Te
Notes on the Foraminifera from Cocos Island.
MILIoLINA PARKERI H. B. Brady.
The examples from the present locality are quite typical. In
its more irregular modifications J. parkeri seems to pass into
M. undosa (Karrer). The nearest locality whence I. parkert was
obtained previously is off Jaffa (Robertson). It is common
outside the atoll, and occurs more sparingly in the lagoon. At
Funafuti it was found close to the inner mar gin of the lagoon.
MILIOLINA LINNZANA (d’Orbigny).
Among the striate forms of MZiliolina occurring at Cocos Island
the above species is worthy of remark. It is very common on the
Jagoon side, and the specimens are of large size and very typical.
On the outer reef it is extremely rare, Tt does not seem to have
been recorded from any dredgings previously obtained from the
Tndian Ocean, but is common in many coral-sands of the Pacific
Ocean and the West Indies.
MILIOLINA FUNAFUTIENSIS Chapman.
This species, which was lately described from the lagoon of
Funafuti (Ellice Islands, Pacific), occurs here in the dredgings
from the outer side of the reef.
HapponiA minor Chapman.
This is another species, occurring not unfrequently, which was
first described from Funafuti.
PAVONINA FLABELLIFORMIS d’Orbigny.
This handsome species was for a long time after its original
description by d’Orbigny almost unknown to rhizopodists, bus it
has of late years been recorded from several localities in the
Pacific and Indian Oceans. It is noted from this particular area
for the first time, and it is worthy of remark that it is found in
greater abundance in the lagoon,
232 MR. F, CHAPMAN ON FORAMINIFERA AND [ Mar. 18,
GAUDRYINA BACCATA Schwager.
Some very fine specimens of this peculiar and redundant form
occur in the lagoon at Cocos Island. It was not met with at all
on the seaward face. The previous occurrences were noted mainly
from deep water ; and in this connection it may be remarked that
it is not uncommon to find certain forms of Foraminifera in-
habiting lagoons as well as deep water, but not in intermediate
conditions of depth.
SPIRILLINA TUBERCULO-LIMBATA Chapman.
This is a form lately found at Funafuti, which is of peculiar
interest on account of its characters partaking of two of Brady’s
species, namely, S. ‘tuberculata and S. limbata. It was found
Cee sparingly at Cocos Island both inside and outside the
agoon.
GYPSINA INHERENS Schultze sp.
The specimens of the above form from Cocos Island are
remarkable for their deep rose-colour, derived presumably from
the sarcode of the animal. This colour was a distinctive feature
of Schultze’s original specimens. The fact of the test being so
strongly tinted points to their fresh condition when collected, for
they seem to lose it very easily, judging from the rarity of its
occurrence. G'. inherens here prefers the quieter water of the
lagoon, but it is also found outside.
PoOLYTREMA MINIACEUM (Pallas), var. ALBA Carter,
A noteworthy feature of these gatherings was the extraordinary
abundance of the white variety of the well-known Polytrema
miniaceum. It seems restricted in its occurrence to the lagoon
deposits.
: Foraminifera from Cocos Island.
Species. Seaward face. | Lagoon.
A ai |
1. | Nubecularia bradyi Millett ..............0...) 0 ss | Vt: |
2. | Biloculina oblonga V@Orb. ..............-- V.0e | scone: |
3. a GLO GL CACLO LD serene cree |e | Welk
4. is depressa W@Orb. ......... 22.0065: v.r. Ne LIS Ee |
5. | Spiroloculina nitida @Orb. ................. v.r. Waits
6. » Uf RU AUER psanesenas Socoe reece V.Cc. | c.
de ae impressa Terg. .............- Vine sh dae
8. | Miliolina circularis (Born) ................. Wil. STAY Be AE
gh. 95 . var. sublineata
Brady: nme ceed | pussoee Ware
10. ss subrotunda (Montagu) .........J 0... r.
11. 3 labiosa (d?Orb:)* ye... oe cae v.r. Soaheasnset vet
12. - tricarinata (d’Orb.), var. ter-
quemiana Brady ............... C. ee wee
118}. se cuvieriana (d’Orb.)) .......-.-:25--| we | if |
14. = SUSUGMES) BVA Ya sss soe sn sdysetesee v.I. Bak
15. 33 boueana (d Orb.) ...............-.. v.I. | f.
16. ” Sunafutiensis Chapman ......... V.1. Ne etecicess
17. 55 linneana (d’Orb.) ............ v.r. VC
f., frequent; ©, common: v.e., very common; r., Tare; V.r., Very rare.
1902.] OSTRACODA FROM COCOS KEELING ATOLL. 233
Foraminifera from Cocos Island (continued).
Species. | Seaward face. Lagoon.
] |
18. | Wiliolina ferussaci (VOrD.).................. V.C. | 75
19: 5 bicornis (W.& J.) ..........000.. @ v.r.
20. 5 JOLASAPO, HENCNY soc spog0nndonpenbone c. fs
21. 4 undosa (Karrer) Gao ee eee
22. 5 seminulum (Linné) ............... f. f.
23. be polygon (CeOxb:)een-ca-pacs cells) ese Cc:
24., a igmaculis)| (Gi@rbs)) 2.2.2... .2.0.----| a PAIRS N Savon
25. 53 oblonga (Montagu) ............... | v.r. | ve Cee
26. 3 transversestriata Brady.........| V.r. IR} Dak eae
27. | Planispirina exigua Brady ..................| Watiires) (ripestikcaseccs
28. | Vertebralina striata d’Orb................... fs mf,
| 29. | Hauerina compressa Orb. ..........005.026:) anes | V.T.
30. | Peneroplis pertusus (Forsk§l) .............. | v.C. | V.c.
31. 7s CUPOAHUPOES (BENIN) soocsoevss0dece|| — gacone | v.r
32. (Monalysidium) Ry TICE
(Lam.) .. ats fe arenas ea cess ere
33. 3 (IL) sollasi Chapman sal fe |e) ER ae Exe
~ 384. | Orbitolites marginalis (Lam.) .. ements V.C. | Wh
35. i duplex Carpenter Sectors taeauecer |). RMR ee | v.r.
RO, || Aliaalhioe TAO CHOWAOS sasccnoaponnopaesoseonesl| . cosene | Yr.
37. | Haddonia minor Chapman .................. fe NRE Ui ean
38. | Textularia conica VOrb. ............ 0000.05. VAT tld es Weer
hers One ‘ ignanmen @Orps aye eee | ¢. | fs |
| 40. PS siphonifera Brady ............... | iF it |
41, 53 OGGRREO GOS CAOTADS \ Sonenscocese|| — aeosor | f.
42. | Pavonina flabelliformis VOrb. ............| v.r. | f
43. | Gaudryina baccata Schwager ............... Ye eaeeee | V.C.
44, | Verneuilina spinulosa Reuss ............... V.c. f.
45. | Vaginulina legumen WOrb. .............0000:) sae | V.1.
46. | Sagraina bifrons Brady ..................... Vil. We meee
47. 3 raphanus Parker & Jones ......) ...... e 1
48. | Globigerina bulloides, var. triloba Rss.... V.c. Cc.
49, . helietiaaOrb: Aes: Gassnie|, ) OTe v.T.
50. ; equilateralis JBRNOKT | Sssecodos| V.I. estes
51. . dutertres @Orp. .............-: lec! 9 eee V.r
52. | Spirillina inequalis Brady .. F ex the eH [el data a
53. 55 tuberculo-limbata Chapman ea ri IR
54, > limbata Brady .. | bh Se, vr
55. decorata Brady... Beseaseseases| (i Mun eateees Vir
56. Cymbalopora poeyi (V’Orb. ) coeenedall Wao V.c.
* BY ‘5 tabelleformis Brady . neat v.r. V.C.
58. Bs (LTretomphalus) bulloides |
(d’Orb.) .. pec aaaeeal C. V.Cc.
59. | Discorbina globularis (a Orb. ys See eer Yr. V.C.
60. 3 polystomelloides P. & J. ow...) sees V.C.
61. | Planorbulina larvata P. & J.. sctieel| (pie crass C.
62. acervalis Brady... gee eee OR OLE AGL. he | V.C.
63. | Car penteria balaniformis Gray ............ v.r. jteed he chee
64, | Pulvinulina menardi (dV Orb.) .. nese Yr. v.r.
65. Pe repanda (f &M,) PS claret aie © allo
66. 5 lateralis Ae Pero dceecs| Cc. 1
67. | Rotalia beccarii (L.) . Meng mete VG} 1
68. | Gypsina globulus (Reuss)... sacageie ge v.r. v.r.
69. bs inherens (Schultze) . Layee f. C.
70. | Polytrema miniacewm (Pallas).............. v.I. V.c.
71. var. alba Carter... V.c.
72. Polystomella erispa (L.) . sen i co. 3 f.
73. im macella (F. & M. ie Bs f, v.r.
74. EA subnodosa Miinster.. EEN Ree ee vere
75. | Amphistegina lessoni d’Orb................... V.c. V.c.
76. | Heterostegina depressa d’Orb................ a Te V.C,
Proc. Zoot, Soc.—1902, Vou. I. No. XVI. 16
234 MR. G. A, BOULENGER ON THE [ Mar. 18,
5. Contributions to the Ichthyology of the Congo.—I. On
some new Fishes from the French Congo. By G. AG
BouLEnGgER, F.R.S.
[Received March 1, 1902.]
(Plates XXII.—XXTV.")
The British Museum has recently received from its excellent
correspondent Mr. G. L. Bates a single fish obtained by him in
the Ja River, flowing into the Sanga, an affluent of the right
bank of the Congo, and this fish proves to belong to an un-
described species which requires the establishment of a new genus
of Siluride.
At the same time the Director of the Royal Brussels Museum
has entrusted me, at the request of my friend M. L. Dollo, with
the study of the Congo Fishes preserved in that establishment.
Among these I have found a small series of specimens coming
from the Lukula River (sometimes spelt Likuala), another affluent
of the right bank of the Congo, parallel to the Sanga. This series
contains examples of five species :—Marcusenius sphecodes Sauvage,
Alestes kingsleye Giinther, Auchenoglanis ballayi Sauvage (all
three previously known from the Ogowe only, and therefore new
to the Congo system), and two new forms which are now described
under the names of Labeo lukule and Chilochromis dupontn.
ALLABENCHELYS, g. 0.
Intermediate between Clarias and Clariallabes. Agreeing with
the former in the free border to the eye, with the latter in the
sides of the head being unprotected by bone.
ALLABENCHELYS LONGICAUDA, sp. n. (Plate XXII. figs. 1, 1 a.)
Depth of body 12 times in total length, length of head 6 times,
Head 11 as long as broad, smooth above, the bony casque, in the
middle, only one third the width of the head; postorbital shield
narrow ; supraoccipital process acutely pointed ; a small frontal
fontanelle ; eye very small, its diameter 4 times in length of
snout and 6 times in interorbital width; latter not quite half
length of head; band of premaxillary teeth 5 times as long as
broad ; vomerine teeth conical, in a crescentic band, which, in the
middle, is nearly as broad as the premaxillary band. Nasal
barbel nearly half as long as head; maxillary barbel as long as
head, reaching middle of pectoral spine; outer mandibular barbel
_ 2 length of head, inner 3. Gill-rakers moderately long, 12 on
anterior arch. Clavicles hidden under the skin. Dorsal fin with
80 rays, anal with 60, both narrowly separated from the caudal ;
the distance between the origin of the dorsal and the occipital
1 For explanation of the Plates, see p. 237,
SWULSIAL (Gls Wy Eh) G WC DIINO SUSITEUET INUEIELY IW" IL
‘dun soig usequrpy TEA? PP F22SNe a.
TXX Id 11°“ C061 S Za
‘HW INMNT O#aV'T fee ee
dar Soag uta Qeeyy WTI TEP ararg “Ld
eat
PCs te los COCs Zc
‘duit: sorg ura quip ILNOdNd SINOYHDOTIHO HIE) LOS) SUES CSL
INES el EA OGL SEI
teak
fe)
1902. | ICHTHYOLOGY OF THE CONGO. 235
process 2 the length of the head. Pectoral half the length of the
head; spine smooth, 2 the length of the fin. Ventrals small,
twice as distant from the root of the caudal as from the end of
the snout. Caudal 2 the length of the head. Dark olive-brown
above, whitish beneath; vertical fins dark, anal edged with white
red 2).
ae length 210 millim.
A single specimen was obtained by Mr. G. L. Bates in the
Ja River, French Congo, 250 miles from the coast.
The nearest ally of this new fish is Clariallabes melas Blegr.,
from-the Lower Congo, which differs, apart from the generic
character of the absence of a free border to the eye, in the longer
head, the more numerous rays to the dorsal and anal fins, which
unite with the base of the caudal, and the presence of serrations
on both sides of the pectoral spine. Clariallabes melas has never
been figured; the upper surface of the head and of the anterior
part of the body is represented on Pl. XXII. fig. 2, for comparison
with Allabenchelys longicauda.
LABEO LUKUL4, sp. n. (Plate XXIII.)
Body compressed, its depth nearly 4 times in total length;
length of head 44 times in total length. Head once and a half
as long as broad; snout obtusely pointed, strongly projecting
beyond the mouth, covered with large nuptial tubercles ; eye
supero-lateral, in the second half of the head, its diameter 6
times in length of head, 23 in width of interorbital region, which
is flat; width of mouth, with folded lips, half length of head ;
rostral flap and anterior border of lip not denticulated ; posterior
border of lip denticulated; inner surface of lp with numerous
feeble, transverse plice; a minute barbel, 2 the diameter of the
eye, hidden in the folds at the sides of the mouth. Dorsal ITT 10,
with notched upper border; the longest ray equals the length of
the head.and twice that of the last ; fin a little nearer the root of
the caudal than the end of the snout. Anal II 5; longest ray
2 length of head. Pectoral falcate, as long as head, not reaching
base of ventral. Ventral reaching vent, its first ray falling under
the seventh (fourth branched) ray of the dorsal. Caudal deeply
forked, with pointed lobes. Caudal peduncle once and a half as
long as deep. Scales 352 ; 4 series of scales between the lateral
line and the root of the ventral ; 12 scales round the caudal
peduncle. Dark olive, belly whitish.
Total length 250 millim.
A single specimen from the Lukula River, preserved in the
Royal Natural History Museum, Brussels. This species is to be
placed near L. macrostomus, L. greenti, and L. nasus, from all three
of which it is easily distinguished by the number of scales round
the caudal peduncle—12 instead of 16 or 18; in this character
agreeing with Z. parvus, which differs in the shorter, less
prominent snout, the shorter caudal peduncle, and one series of
16*
236 ON THE ICHTHYOLOGY OF THE CONGO. [ Mar. 18,
scales less both above and below the lateral line. The numbers of
scales are as follows in the six Congo species with notched dorsal
fin with 10 or 11 branched rays, supero-lateral eyes, and a single
barbel on each side :—
L. falcifer Blgr.—S8q. 39 a 5 between L. 1. and V., 20 round
caud. ped. ;
L. macrostomus Blgr.Sq. 388-3975, 4 between L. 1. and V.,
- 16-18 round caud. ped.
L. greenit Blgr.—Sq. 37-38 ° 4 between L.1. and V., 16 round
caud. ped.
LL. nasus Bler.—8q. 38-39 4 between L.1. and V., 16 round
caud. ped.
L. lukule Blgr.—Sq. 35 4 between L.1. and V., 12 round
caud. ped.
ii poke Bee 33-35 =, 3 between L. 1. and V., 12
round caud. ped.
CHILOCHROMIS, g. n.
Body moderately elongate; scales cycloid. Jaws with very
broad bands of bristle-like movable teeth with club-shaped inbent
crowns; rami of lower jaw approximated, spatulate in front,
connected with the upper jaw by a broad, thin lip; maxillary
concealed under the preorbital. Dorsal with 17 spines, anal
with 3.
This remarkable new genus approaches Petrochromis Bler., but
differs from it in the narrower lower jaw and in the teeth being
simply club-shaped, instead of bi- or tricuspid. I have much
pleasure in naming the species in honour of the eminent Director
of the Brussels Museum, one of the pioneers in the geological
exploration of the Congo Basin.
CHILOCHROMIS DUPONTI, sp. n. (Plate XXIV.)
Depth of body 23 times in total length, length of head 32 times.
Snout rounded, with arched profile, a little longer than the
diameter of the eye, which is contained 34 times in the length of
the head and 14 in the interorbital width; mouth extending to
below the nostril; teeth very numerous, with reddish-brown
crowns, those of the upper jaw in 8 or 9 transverse series, those of
the lower jaw forming two spoon-shaped groups; the inner teeth
smaller than the outer; 3 or 4 series of scales on the cheek;
large scales on the opercle. Giull-rakers short and slender, 15 on
lower part of anterior arch. Dorsal XVII 10; last spine longest,
2 length of head; middle soft rays longer than the head. Pec-
toral acutely pointed, as long as the head. Ventral not reaching
the vent. Anal III 8; third spine longest, a little shorter than
longest dorsal; middle soft rays # length of head. Caudal fin
feebly emarginate. Caudal peduncle as long as deep. Scales
33 = lat. 1. =. Uniform olive-brown above, yellowish beneath ;
fins greyish.
PG, Sr NO2 roll Ik ee ZOay
J. Smit del.et hth, MinternBros.imp.
CERCOPITHECUS OTOLEUCUS
Se
1902. ] THE SECRETARY ON ADDITIONS TO THE MENAGERIE, 23h
Total length 220 millim.
A single specimen from the Lukula River, preserved in the
Royal Natural History Museum, Brussels.
I avail myself of this opportunity to propose the name of
Pelmatochromis polyodon for the fish from Monsembe which I
have recently described (Ann. Mus. Congo, Zool. ii. p. 53) as
P. teniatus, having overlooked the fact that the same name had
been previously bestowed on a species from Nigeria.
EXPLANATION OF THE PLATES.
PratE XXII.
Fig. 1. Allabenchelys longicauda, p. 234, 3 nat. size.
la ; Upper surface of head, } nat. size.
“O) 3” 5) =
2. Clariallabes melas, p. 235, 3 nat. size.
Prare XXIII.
Labeo lukule, p. 235, with view of open mouth, § nat. size.
Puate XXIV.
Chilochromis duponti, p. 236, with view of open mouth, + uat. size. _
April 15, 1902.
Prof. G. B. Howrs, LL.D., F.R.S., Vice-President,
in the Chair.
The Secretary read the following report on the additions made
to the Society's Menagerie in March 1902 :—
The registered additions to the Society’s Menagerie during the
month of March were 146 in number. Of these 38 were acquired
by presentation, 18 by purchase, 3 were born in the Gardens, and
87 were received on deposit. The total number of departures
during the same period, by death and removals, was 143.
Amongst the additions attention may be specially directed
1@ B=
i. A Monkey of the genus Cercopithecus, procured by Major
Delmé-Radcliffe in the Latuka Mountains, about a hundred miles
east of the Upper Nile in Northern Uganda, and presented to the
Society on March Ist. This Monkey appears to belong to a new
species allied to Cercopithecus lewcampyx, but easily distinguish-
able by the white ear-tufts and grey back. I propose to name
it Delmé-Radeliffe’s Monkey (Cercopithecus otoleucus). It may
shortly be described as follows :—
CERCOPITHECUS OTOLEUCUS. (Plate XXV.)
Above fuliginous, back more or less grizzled with pale fulvous,
head above black; frontal line white, with hairs rather elongated ;
238 _ ON SPECIMENS OF FOSSIL BONES FROM cyPRuUS. [Apr. 15,
ears blackish, with a conspicuous patch of white hairs in the lower
part of the conch; sides of face grizzled like the back, but more
greenish ; nose blackish, chin whitish ; limbs and tail black; belly
and underparts pale whitish grey: whole length of body about
13 in.; tail 17 in.
Hab. Forests of Latuka Mountains, Northern Uganda.
Obs. Closely allied to C. leucampyx of West Africa, but distin-
guished by its white ear-patches, blacker head, greyer back, and
much paler colour beneath.
[P.S. July 1st.—Herr Oscar Neumann, who has examined this
Monkey, is of opinion that it is nearly allied to, if not identical
with, Cercopithecus stuhlmanni of Matschie (Sitzsb. Ges. naturf.
Fr. Berlin, 1893, p. 225). ,This is possibly the case, but the
description does not quite agree with our specimen. |
2. A Panda (dlurus fulgens), from Northern India, obtained
by purchase on March 4th. ‘This scarce animal, which is the
third specimen received by the Society, was in a weak state on
arrival and unfortunately did not live long.
3. Another collection of ten Indian Birds, presented by Mr. E.
W. Harper, F.Z.S., all belonging to species new to the Collection.
Amongst them the Stork-billed Kingfisher (Pelargopsis gurial)
and the Mountain-Thrush (Oreocincla dawma) are particularly
interesting.
Prof. Bell, F.Z.8., exhibited two arms of an injured Starfish of
the genus Lwidia, from the west coast of Ireland, which had
undergone repair at their free ends. These regenerated parts
were unlike the rest of the arm, and had a striking though not
exact resemblance to the free ends of the arms of an Astropecten.
Dr. Forsyth Major, F.Z.S8., exhibited some selected specimens
from a collection of fossil bones recently received by the Natural
History Museum from Cyprus, where they had been discovered in
caves by Miss Dorothy M. A. Bate, who started last year for that
island with the express purpose of discovering and exploring
ossiferous caverns.
The remains proved to be those of a pigmy Hippopotamus,
about half the size of a middle-sized Hippopotamus amphibius,
and could not be distinguished from Cuvier’s “ Petit Hippopotame
fossile” (H. minutus Blainv.), which was smaller than the so-
ealled ‘ H. minutus” of Malta and otherwise different.- Cuvier’s
description had been based on scanty remains in the Paris Museum
and from private collections in Bordeaux and Brussels, all of them
without any record of their origin, but which had ultimately (Oss.
Foss. 4th ed. ii. p. 490, 1834) been supposed to come from a place,
never identified before nor after, between Dax and Tartas in the
South of France. Dr. Forsyth Major now suggested that the
fossils described by Cuvier were, in reality, from Cyprus also.
1902.] ON THE ANATOMY OF THE CONDOR. 239
The fossils exhibited showed affinities, on the one hand, with -
the pigmy Hippopotamus of Western Africa, Cheropsis liberi-
ensis; on the other, with some remains from the Lower Pliocene
of Casino (Italy). They were considered by the exhibitor as a
further illustration of the assumption that many of the Pleistocene
Mammals of the Mediterranean Islands were the little modified
survivors of Tertiary forms from the adjoining continents, from
which the islands had been severed during that period.
The following papers were read :—
1. On the Windpipe and the Heart of the Condor. By
Frank K. Bepparp, M.A., F.R.S., Vice-Secretary and
Prosector of the Society.
| Received March 12, 1902. ]
(Text-figures 29-32.)
The generalities in the structure of the windpipe of the Condor
are pretty well known, and have been so for long, though the
information as given is not always exact. There has not been,
however, so far as I am aware, a detailed comparison of that organ
in the two sexes. As sexual differences in the windpipe are to
be found at least in Sarcorhamphus gryphus, | have thought it
worth while to draw up an account of the matter. In the female
Sarcorhamphus gryphus the two bronchi end, as I described and
figured them some years since’, in a membranous tract of some
length ; the cartilaginous rings of the bronchi in fact cease to
exist some way before the bronchi plunge into the lung-substance.
This membranous tract of each bronchus is enveloped and com-
pletely covered by a layer of muscle, which is prolonged into
several strands of muscle tying the bronchus down to the mem-
branous surface of the lung. The figure of the windpipe in the
female Condor illustrating my account of it was drawn from the
recently dead specimen, and is, I believe, quite accurate. I have
examined also the windpipe of a female example of the second
species of the genus Sarcorhamphus, viz. S. equatorialis, which I
had preserved at the time of the death of this specimen. The
end of each bronchus is, in precisely the same way as in S. gryphus,
covered with a sheet of muscle. I do not give here a detailed
account of the arrangement of the muscular tags proceeding from
this sheath of muscle and tying down the bronchus to the lung-
surface, since they appear to be, as far as I can judge, identical
with the arrangements to be seen in Sarcorhamphus gryphus.
I have quite lately had the opportunity, through the death of
one of these birds, of examining the windpipe in the male Sarco-
1 “Notes on the Anatomy of the Condor,” P. Z.8. 1890, p. 142.
240 _ MR. F. E, BEDDARD ON THE [Apr. 15,
rhamphus gryphus. I exhibit drawings of the windpipe (text-
figs. 29, 30), which are quite accurate, and illustrate the various
features to which I desire to direct attention. It will be seen
Text-fig. 29.
en
Lower end of windpipe of male Sarcorhamphus gryphus, front view.
M, extrinsic muscles ; T, muscular tag at end of bronchus.
that each bronchus ends in the male bird, as it does in the
female, in an entirely membranous section of considerable length
in proportion to the entire bronchus. The proportions are rightly
1902. ] ANATOMY OF THE CONDOR. 241
shown in the drawings to which I have called attention. Here,
however, the resemblance ends. For in the male bird I can find
no trace visible to the naked eye of the muscular sheath which
covers this part of the bronchus in the hen bird. I have care-
fully looked for these muscles both in the fresh windpipe and
after it had been preserved in alcohol. There is no doubt in my
mind that the difference indicated does really exist. Moreover,
the rather abundant slips of muscle, which, in the hen bird, tie
down the end of the bronchus to the membranous surface of the
Text-fig. 30.
Lower end of windpipe of male Sarcorhamphus gryphus, back view.
M.T, membrana tympaniformis.
lung, are only just recognizable in the cock bird. I found but
one tiny slip (T in text-fig. 29) representing this very striking
feature of the windpipe of the female. This required careful
looking for ; there is nothing to strike the eye forcibly.
It appears, therefore, that in Sarcorhamphus gryphus certainly, -
and in Sarcorhamphus cequatorialis probably, there is a marked
difference between the sexes in the conformation of the syrinx ;
and that, in the second place, the more complicated organ is that
of the female, and not of the male sex. These birds have no
242 MR. F. E, BEDDARD ON THE [Apr. 15,
proper voice, but only “hiss like a reptile.” Whether the
musculature of the lower part of the bronchi in the females pro-
duces any difference in the sounds uttered, I am not aware.
This is the principal fact in the structure of the windpipe
of Sarcorhamphus to which I have desired to call attention.
There is, however, another matter to which I did not give any
attention in my description of the windpipe of the female Condor.
Prof, Fiirbringer has remarked that “ Die... . Cathartidee heben
sich durch vollkommene bronchiale Ringe besonders hervor.”* I
have briefly referred to the fact” that in the members of this
group there is, at least sometimes, an imperfectly formed tracheo-
bronchial syrinx—imperfect in the fact that there is no great modi-
fication of the rings at the bifurcation, but suggesting a tracheo-
bronchial syrinx in the incompleteness of the bronchial rings,
which are indeed semi-rings. In Sarcorhamphus gryphus this
incompleteness of the bronchial rings is very plain, as the drawing
submitted herewith (text-fig. 30, p. 241) shows. Moreover, the
membranous space lying between the approximated ends of the
bronchial semi-rings on the dorsal aspect of the windpipe is
prolonged upwards for a considerable distance along the trachea,
gradually diminishing until the tracheal rings, at first incomplete,
become fully complete rings passing without a break right round
the windpipe. There is even—at least I so interpret it—a slight
suggestion of a pessulus; this is in the shape of a small piece of
cartilage, again divided into two, which lies at the point of
bifureation of the trachea into the two bronchi.
I have had an opportunity of comparing the windpipe of the
Condor with that of the American Vulture, its near ally, Cathartes
atratus, of which several specimens have recently died in the
Society’s Gardens. These examples were of both sexes, and I
have not been able to note any sexual difference such as charac-
terizes the Condor. In Cathartes, moreover, the bronchus,
although it does end in a short membranous tract, is not invested
with muscle as in the female Sarcorhamphus. In fact, the
appearance of the bronchi at their termination in the lung is
much like that of the male Sarcorhamphus. Furthermore, in
each case that I examined, three muscles attached at one end to
the ribs were inserted on to the surface of the lung in the vicinity
of the entrance of each bronchus. It seems to be reasonable to
compare these muscles, which are of course the usual lung-muscles
(‘ diaphragm” of some authors), to those found in most birds
arising from the ribsand implanted upon the membrane covering
the lungs. But in addition to this comparison, it seems also
possible to compare them especially with the “tags” of muscle
already described in the Condor as connected with—or, indeed,
arising from—the sheet of muscle covering the membranous
termination of the bronchus. In Cathartes, however, all trace of
an attachment to the bronchus itself was lost; the muscles are
1 Untersuchungen z. Morph. u. Syst. d. Vogel, Amsterdam, 1888, p. 1086.
2 The Structure and Classification of Birds, London, 1898, p. 482.
1902. ] ANATOMY OF THE CONDOR. 243
plainly, as in other birds, inserted on to the surface of the
membrane covering the lung. Nevertheless, it appears to me
that possibly the conditions obtaining in these two kinds of birds
may give a clue to the origin of a portion of this musculature of
the lung. I have suggested already, in my former paper upon
the Condor, that the sheet of muscle enwrapping the end of the
bronchus may be the remnant of an intrinsic syringeal muscle
altered in function in correspondence with the degeneration of
the syrinx itself. A second stage is seen in the male Sarco-
rhamphus, where the muscle is reduced to the one or two “ tags”
which tie down the end of the bronchus to the lung-surface.
The final stage is shown in Cathartes, in which birds in both
sexes the “tags” of muscle are present and well developed, but
have entirely lost all special relation to the end of the bronchus.
This, however, is at present a suggestion for the origin of those
muscles, the nature and distribution of which require, and have
not yet had, detailed attention in many groups of birds.
The heart of the Condor has been dealt with by Gegenbaur '
and by myself *. We have both recorded the occurrence of traces
of the septal flap of the right auriculo-ventricular valve, which is
for the most part missing in Birds. I therefore examined with
particular interest the heart of the male Sarcorhamphus gryphus,
to the windpipe of which I have directed attention above.
Gegenbaur found in a heart examined by himself “a fold....
which is formed by a thickening of the endocardium”; this fold
was found to run backwards ‘from the anterior origin of the
muscular valve.” It is not altogether easy to follow the description
given by Gegenbaur, since it is unaccompanied with any drawings.
I take it, however, that what Gegenbaur saw in the heart studied
by him was a prolongation of that part of the valve, arising to
the left of the great papillary muscle, tying the valve to the
free non-septal wall of the ventricle: to the left, that is to say,
when the right ventricle is opened and looked at from in front.
Now I have already brought forward reasons for considering that
this part of the valve, which appears occasionally to be rather
membranous in constitution and is always of thinner texture
than the larger half of the valve, is the equivalent of the septal
half of the valve in the Crocodile’s heart *. A further study of the
Crocodile’s heart led Dr. Chalmers Mitchell and myself to the
same conclusion. What Prof. Gegenbaur therefore has been able
to place on record is a still greater extension of this septal half
of the right auriculo-ventricular valve. In a vanishing structure
such fluctuations are generally met with.
In the corresponding valve of the Monotreme heart, Prof.
Lankester found considerable variations in the amount of the
septal half of the valve which was present; and I do not doubt
that careful measurements would prove the same thing for the
1 “Zur vergleichenden Anatomie des Herzens,” Jen. Zeitschr. ii. 1866, p. 365.
2 P.Z.S. 1890, p. 144.
3 “On the Structure of the Heart of the Alligator,” P. Z.S. 1895, p. 348.
244 MR. F. E. BEDDARD ON THE [ Apr. 15,
rudimentary septal flap of other birds. The traces of the septal
flap, other than this definite piece of that flap noted by myself in
the Condor’s heart, consisted in ‘4a series of tiny yellowish spots
and vesicles a little way from the posterior margin of the atrio-
ventricular orifice, which formed a line occupying a_ position
identical with that which would be occupied by a septal part of
the valve if it were present.” These structures, possibly patho-
logical, seemed to me and still seem, to be possibly a reminiscence
of that half of the valve. Apart, however, from this pathological
and thus more questionable state of affairs, we have Gegenbaur’s
positive assertions. In the heart which I have most recently
Text-fig. 31.
Heart of Sarcorhamphus gryphus opened so as to display the interior of the
right ventricle. ;
A, cut end of papillary muscle tying valve to septal wall of heart ;
B, opposite end of the same; C, rudiment of septal flap.
examined there were no traces, that I could discover, of an exten-
sion of the top half of the septal flap; but, on the other hand, as
is shown in the drawing submitted herewith (text-fig. 31), a con-
siderable piece of muscular tissue extended from the lower end
of the invariably present portion of the valve in the direction of
the rudimentary slip at its other end. A line joining the letters
A and B in the drawing would make a complete septal half to
this valve. It cannot, I think, be doubted that this structure is
a rudiment of the chiefly missing septal half of the valve ; and if
(1902. ] ANATOMY OF THE CONDOR. ~ 245
the two hearts, this one and the heart described by Prof. Gegen-
baur, were to be reproduced in a composite drawing, we should in
all probability see a bird’s heart with a right auriculo-ventricular
valve as complete as is that of the Crocodile. It may be noted,
moreover, that in the Crocodile’s heart (See the figures illustrating
the paper by myself and that by Dr. Mitchell and myself) the
portion of the septal half of the valve which is nearest to the half
of the valve attached to the free wall of the right ventricle is
entirely muscular, the fibrous portion of the valve lying more to
the right. The comparison therefore becomes so far more exact.
A still further reduction of this already reduced representative of
the lower half of the septal flap of the valve might result in such
small muscular pillars, arising from the ventricular septum and
connecting together the two walls of the ventricle such as occur
in various birds, and which I have especially called attention to
in the heart of Chunga in my paper already quoted.
In addition to the heart of the Condor, I have had recently the
opportunity of examining the heart of another bird which shows
Text-fig. 32.
Heart of Scythrops nove-hollandie cut open so as to display interior of
right ventricle.
A, papillary muscle; B, tendinous seam extending from the same ; C, valve.
some persistent traces, as I regard certain structures to be
described immediately, of the missing half of the right auriculo-
ventricular valve. The accompanying sketch (text-fig. 32) is a
drawing of the heart of a Cuckoo (Scythrops nove-hollandic),
from which, as in the case of the Condor’s heart just described,
the apex has been removed. The right ventricle is cut and
reflected, and the complete half of the valve shown without
further cutting. From the left half of the valve, which is attached
to the interventricular wall, a white seam runs down the ventri-
cular wall in the same direction as the piece of valve from which
it originates. Unfortunately, as will be seen from an inspection
of the sketch, the removal of the apex of the heart has destroyed
the actual ending of the seam, It could not end, however, very
246 ON THE ANATOMY OF THE CONDOR. [ Apr. 15,
far from the end of the complete half of the valve. The seam, as
I call it, has hardly a muscular or tendinous appearance. It is
not perceptibly raised above the muscle which forms the wall of
the heart; it is only conspicuous through its decided white
colour. This description, I should observe, applies to the heart
after preservation in spirit. I received the whole bird from
Australia in spirit some years since. But the seam is so marked,
that I cannot but think that it would have been as obvious in
the fresh heart as it is in that preserved in spirit. The appear-
ance of the seam, in fact, suggests a thickening of the lining-
membrane of the heart, the endocardium. It just runs on to
the commencement of the left-hand piece of the muscular existing
valve. Now it appears to me to be fair to construe this structure
as a remnant of the otherwise chiefly missing septal flap of the
atrio-ventricular valve. It may be admitted that its course is
straighter than such a flap had when fully developed. But with
rudimentary structures, alterations of one kind or another not
related to their former functions are not uncommon. I do not, in
fact, think that the length of the seam is against my interpreta-
tion of its nature. As to the possibility that it is a thickening of
the endocardium, it seems to me that it is then very comparable
to the “fold” described by Gegenbaur, “‘ which is formed by a
thickening of the endocardium.” And Gegenbaur adds to the
description that the fold in question arises “from the anterior
origin of the muscular valve on the septum ventriculorum,” which
is precisely the origin that the seam described here by myself has.
Gegenbaur’s fold, however, runs “obliquely backwards and down-
wards,” so that its position as a rudiment is more in accord with
that interpretation. A final point is of some little interest. It
is or has been believed that ontogenetically as well as phylo-
genetically the muscular or tendinous valves of higher vertebrates
are first formed as simple thickenings of the endocardium, later
invaded by muscle which itself later on is converted into tendon
(as in higher mammals). The return, so to speak, of this rudi-
ment of the septal half of the valve to its very earliest condition
is worth emphasizing.
The facts that have just been dealt with raise another inter-
esting question. At one time the descent of Birds from some
Dinosaurian form was widely believed in; later this view lost
some ground, until quite recently Prof. Osborn has recommended
its serious reconsideration mainly on the grounds of the discovery
of a fourth toe bent backwards, which has been shown to exist in
the Dinosauria. This and some other features have added not a
little to the bird-like characteristics of that group of Reptiles.
On the other hand, there have not been wanting those who would
assign the origin of Birds to a lower type of Reptile. The nature
of the heart-valves seems to me to throw some light upon the
question. At first sight, the arrangement of the auriculo-
ventricular valves in the bird is more suggestive of the same
valves in the tortoise than in the crocodile, the latter repre-
uae | %
0 Gis
: Taue
ueny
PAsS IO 2-vol I ereoaie
F.Pickard-Cambridge del.et lith. West,Newman imp.
SPIDERS OF THE GENUS LATRODECTUS.
Bee
ro
1, 1S, IS Oa rroll I. led POOvs,
E Pickard-Cambridge del.et lith. West,Newman imp.
SPIDERS OF THE GENUS LATRODECTUS.
1902. ] ON THE SPIDERS OF THE GENUS LATRODECTUS. Q47
~ senting the type of an highly-organized reptile, and nearer to the
Dinosaurs than any other living type. As in the genus Testudo,
the right auriculo-ventricular valve of the Birds is almost always
a clearly-defined and semi-lunar muscular flap, practically guarding
only one side of the auriculo-ventricular orifice. In Testudo, as in
the bird, the opposite side of the ventricle has a perfectly smooth
surface, without even the rudiment of a ridge to correspond to the
valve which lies on the left side of the auriculo-ventricular
orifice. On the other hand, the left auriculo-ventricular aperture
is guarded by a completely, or nearly completely, encircling valve
in both tortoise and bird. Hardly any change is required to
convert the right auriculo-ventricular valve of the tortoise into
that of the bird. If no rudiments, such as those described in the
present communication, of a septal half to the right auriculo-
ventricular valve had been discovered, it might be reasonable to
dwell upon the striking but really superficial likeness which this
valve in the vast majority of existing birds shows to the corre-
sponding valve in the tortoise. But it seems now to be clear
that the simple valve in the bird’s right ventricle is not the
persistent and simple valve of the tortoise or some lower
reptilian type; but that it has been derived from the reduction
of a more complicated valve such as is possessed by the Crocodiles,
and was very possibly possessed by the Dinosaurs. The evidence
is therefore so far in favour of assigning to the birds an origin
from some highly-developed reptilian type.
2. On the Spiders of the Genus Latrodectus, Walckenaer.
By Freperick Pickarp Campripas, F.Z.S.
[Received February 25, 1902. ]
(Plates KXVI & XXVII.)
ConrTENTS.
I. Introduction, p. 247.
II. List of Species and Sub-species recognized in this paper, p. 252.
III. Alphabetical List of Species described or figured, p. 256.
IV. Explanation of the Plates, p. 261.
I. IntRoDUCTION.
The genus Latrodectus of Walckenaer (Tableau, p. 81, 1805),
of which the type is L. 4-tredecim-guttatus (Rossi), 1790, com-
prises those very interesting Spiders which, under various local
names, have been notorious in all ages and in all regions of the
World where they occur on account of the reputed deadly nature
of their bite.
As to the evidence, there can be no doubt whatever that the
inhabitants of those regions suffer frequently from blood-poisoning
1 For explanation of the Plates, see p. 261,
248 - MR, F, PICKARD CAMBRIDGE ON THE [Apr. 15,
of a very serious nature, and that wherever the variety of Latro-
dectus occurs which is of black coloration with vivid red spots,
particularly at the apex (or tail- end) of the abdomen, the blood-
poisoning is ascribed to the ‘‘sting” of this spider. Of course
the spider has no sting at the tail- end, but people who feel a
sudden wound are not likely to note very carefully which end of
the enemy is responsible for the damage. The tail-end being
briliant red, however, looks full of venom, and hence they have
jumped to the conclusion that the sting lays there. To such an
extent has this belief prevailed that in Guatemala, Mr. Sarg
relates, the natives assert that the spider actually squirts out its
venom at the victim, and if the fluid even touches the skin, an
angry eruption will supervene. The fluid which is thus ejected
is not venom, however, but merely liquid-silk, an adhesive
treacly fluid which the spider squirts at a captive in order to
impede its struggles for liberty. In many species of the family,
Theridiide, to which Latrodectus belongs, the same phenomenon
can be observed.
One may be pardoned for suspecting that the red colour has
given rise to the supposition that the “sting” of these spiders is
extremely venomous; and whether they are the true culprits or
not, suspicion would naturally fall upon them at once amongst
the ignorant and unobservant.
I have not heard that any venomous tendencies have ever
been ascribed to ZL. geometricus, a large grey species, abundant
in houses, outbuildings, and offices, where, if members of the
genus are as bloodthirsty as has been alleged, adults and children
are sure to have been bitten or “stung.” I never once heard
any of the natives accuse this species of stinging, or utter
warnings as to the poisonous nature of its bite, though it was
numerous in the houses throughout the Lower Amazons, and
they were not behindhand in making the most of the dangers of
the forest and the deadly nature of its inhabitants. But, then,
there are no red spots on this species.
Of course it may also be argued that only those species with
red spots are poisonous, and that the red colour is one of nature’s
danger signals; considerations which bring us no further towards
a settlement of the question. e
Mr. Crotch, writing in 1865, referring to the variety of the
common European form found in Hierro, one of the Canary
Islands, says that his entomological enthusiasm was checked by
solemn warnings as to the dangers incurred from the bite of a
certain large black lethiferous Latrodectus malmignatus vav.,
generally causing death unless relieved by timely and internal
doses of human excrement. A little later he remarks that, so
great was the contempt induced by familiarity, that he could not
be restrained from picking up the deadly monster, which, though
tormented in the way presumed to be most provoking to a spider,
persisted in lying inert in his hands, nor could it be induced to
bite by any means.
1902. ] SPIDERS OF THE GENUS LATRODECTUS, 249
The late Dr. Marx, too, tried many experiments on various
animals by injecting the contents of the poison-sac of these spiders,
but without any prejudicial results,
Numberless instances, however, have been quoted by authors
of cases in which human beings, supposed to have been bitten by
members of this genus, have manifested very serious symptoms.
And this is especially the case in connection with the Katipo, the
New Zealand variety. Mr. Urquhart says that his friend
Mr. Robson stated that a man engaged in erecting a lighthouse
was bitten on the neck, and went quite mad for some time.
Mr. Wright, in 1869, gave many cases: an Knglish boy was
bitten as he sat amongst the grass inhabited by a Katipo; in
another case the spider crawled up a boy’s trousers and gave him
a most dangerous wound. Carl Lumholtz, in his book ‘ Amongst
Cannibals,’ says that Latrodectus hasseltii, the Australian form, is
very common and very dangerous to man; a friend of his, bitten
in the leg, was seized with paralysis for three days. On the
third day he had a cold perpsiration, and recovered.
There are said to be two kinds of Katipo—one black without
spots, the other black with red spots; and the latter is regarded
as far the most poisonous.
One might quote instances of these distressing symptoms from
all parts of the globe—North America, Chili, Central America,
Australia, Madagascar, New Zealand, and the Mediterranean ;
and I must refer readers to the cases mentioned in the ‘ Bulletin’
of the U.S. Department of Agriculture, for Jan. 1889 and onwards
(see the list of Literature), for some startling cases of blood-
poisoning, possibly caused by spider-bites, though there is no
evidence that the bites were inflicted by Latrodectus.
But for well-authenticated cases of men, women, children, and
animals manifesting the symptoms of blood-poisoning, cases
which have been watched for days by medical men who may be
presumed to have been competent observers, we must refer to
the encyclopedic work of Dr. Puga Borne, published in 1891, —2,-3.
Here we have stated the age, sex, condition and temperament of
the sufferers, the time of the year, and the condition of the
weather at the time the patient was bitten. Then we have
minute diagnoses of the symptoms, followed by a careful con-
sideration of the various remedies and their effect.
But both in the works referred to before and in the present
case, there is always a noteworthy and important omission. No
mention is made as to what evidence there was that the bite was
actually inflicted by the spiders accused. Dr. Puga Borne states
that a sheep was bitten by five of these spiders, but he does not
say how they were encouraged to bite, how they were held, or
what part of the sheep was bitten. One cannot believe that so
many cases could be quoted by a medical man without he had
actually witnessed the act of biting, but it is an extraordinary
thing that he should not have thought it worth while to tell us
- how the experiments were carried out and how so venomous a
Proc, Zoou. Soc.—1902, Vou. I. No, XVII. Leh
250 MR, F, PICKARD CAMBRIDGE ON THE [ Apr. 15,
creature was held when being applied to the patient! The
mandibles of Zatrodectus are so insignificant that they would,
for instance, scarcely pierce the thick skin of the fingers.
In the face of so much evidence, however, one would not wish
to suggest that the animals whose symptoms are described were
not actually bitten by the spider in the presence of and under the
eye of a competent observer; only one must insist that it is a
great pity that, if so, this is not distinctly stated and the modus
operandi described by Dr. Puga Borne. _ It is so easy to settle the
point once for all, and so silence scepticism. In every case, how-
ever, which has come under my notice, it has always been taken
for granted at the outset that the wound was inflicted by the
spider; and this being the only point on which proof is desirable,
we are given abundant evidence on the character of the sub-
sequent symptoms and every other point except this.
A New Zealand friend of mine assured me that his friend’s dog
had been bitten by a Katipo, and fancied he had proved the case
up to the hilt when he declared that he had seen the dog running
round in circles on the lawn. Certainly, there must have been
something seriously wrong with his friend’s dog, but the lament-
able symptoms described were no proof that a Xatipo was the
cause of the trouble.
One would therefore urge those who happen to be in the
Katipo country to try and get first-hand evidence. It is not
sufficient to see a AXatipo in the neighbourhood of the tragedy,
nor to see one crawling on the victim’s clothes, nor even to find
one crawling under the clothes of the person bitten. If a dozen
healthy boys could be induced for a consideration to allow them-
selves to be bitten under competent observation, the matter
might be settled once and for all—or, failing this, experiments
might be made on mice, though one cannot suggest how the
spiders are to be induced to bite in either case.
These reports of the symptoms manifested in cases coming in
for treatment suffering from the supposed bite of the spider are
of no value whatever as evidence as to what was the originating
cause of those symptoms.
It may be interesting to gather together the various names
under which the members of this genus are known in the different
countries where they enjoy such evil reputation.
In New Zealand it is known as the Xatipo; in the Philippines
as the Lawalawa ; in Mexico and Central America as the Arana
capulina, because of the resemblance which the abdomen bears to
the fruit of the Capollin cherry (Cerasus capollinus) ; in Guatemala
as the Casampulga; in Madagascar as the Menavoudi; in Chili
as the Pallu or Guina; in Italy as the Malmignatte; in the
Russian Steppes as the Karakurt; in Bolivia as the Mico; in
Peru as the Zucacha; and in the Antilles as the Arata naranja.
I must here express my thanks to M. EH. Simon, Prof. Kul-
ezynski, Mr. H. R. Hogg, and Mr. R. I. Pocock for kindly allowing
me to examine specimens of the genus; and I am also indebted
1902.] SPIDERS OF THE GENUS LATRODECTUS. 251
to Mr. R. Jenery-Shee, an expert in European languages, for
kindly looking thr ough Dr. Puga Borne’s voluminous work in
Spanish, in case I should have missed the paragraph which might
have contained the evidence I required.
Of the forty-three described species referred to this genus, I am
able to recognize six only as distinct, and perhaps eight as sub-
species. Of the former, L. hystrix, geometricus, pallidus, tredecim-
guttatus, and mactans are probably good species. . As to the others,
it is very difficult at present to take up any decided position with
regard to them, as must always be the case where we have under
consideration for ‘ms which are actually, at the time of observation,
undergoing those processes of differentiation, under the influence of
individual variability combined with that of physical surroundings,
which, in these early stages, have not yet brought about any
definite structural difference, or even any variation in the more
superficial characters of colour-pattern, which can be considered in
any way constant and exclusive.
In the subjoined table will be found the characters by which
the most distinct of the species and sub-species of the female sex
can best be recognized. The males are not sufficiently well known
to enable one to tabulate their characters.
Females.
A. Integuments clothed with small acanthoid spines
and short stiff black spines ............::2-c0cee eee seeees hystrix Simon.
B. Integuments clothed with fine short acanthoid
spines and longer bristles, or with fine hairs only.
I. Central anterior eyes eae larger than the
Hlaterals| te ete nee tee cy ue ROCOMCERICUS\G, LieKkorns
II. Central anterior eyes not _ oer fee the oe
1. Integuments almost glabrous. Latero-ventral
area clothed with acanthoid spmes only.
Abdomen entirely creamy-white, with the
black impressed muscular scars very con-
spicuous, and sides slashed with brownish
yellow ......... . pallidus O. P. Cambridge.
a. Hyes of anterior ‘row, ‘asa yule, equidistant}.
a‘, Size much larger, length from 12-14 mm.
Abdomen either ‘entirely black or brown,
without any red spots or with a single
square or elongate-oval red spot above the
anal tubercle; or with a narrow central
dorsal red stripe, broken into two round
spots anteriorly, and with, or without, two
oblique lateral red stripes. Ventral spot
dumbbell-shaped, without a decided dark
spotintthenmmiddletgee 5. v.24 soak ess naee toes mactans Fabricius.
61. Size much smaller, length 7 millim. Ab-
domen rich brown, with three irregular
transverse crimson cinctures (very variable
in exact form however) and a central
posterior crimson band. Ventral spot
oblong-oval, with a decided dark spot or {
blotch i qiihe middle. fo Sst Aaees eon che ees curacaviensis Miller.
geographicus Hasselt.
1 Characters drawn from the eye-formula are not reliable; they vary very much,
even amongst examples from the same district. ie
252 MR. F, PICKARD CAMBRIDGE ON THE [ Apr. 15,
b. Eyes of anterior row not equidistant, centrals
nearer together than to the laterals !.
[ Note.—The following three species are, so far
as I am able to judge from the material at
hand, all of one fundamental form, namely,
tredecim-guttatus Fabr., but they fall
into certain groups more or less limited
by locality. Subjoined are the characters
of extreme examples. |
a*, Legs of first pair longer in proportion ;
tibia i. at least one-fourth longer than
the carapace.
1. Abdomen either entirely black or brown, or
with a central red band more or less broken
up into distinct spots and three or more oval-
elongate lateral spots. Ventral spot either
absent or represented by a transverse band
immediately below the genital rima, and often
one above the spinners; probably also con-
fluent, forming a larger spot .....
2. Abdomen black, with an oblong-oval, “central,
posterior apical red spot, with three ver y small
white spots on each side. Ventral spot usually
represented by a transverse bar below the
genital rima
3. Abdomen black, with a single central ‘longi-
tudinal red band, anteriorly “either constricted
or broken off to form a separate spot. Ventral
spot either absent or with one or two trans-
verse spots, or with a large ES
blotch . hasseltii Thorell.
b2, Legs of first pair ‘shorter in pr -oportion ;
tibia i. not longer than the carapace.
Spider usually Gmialler Aean ceseliec crete katipo Powell.
tredecim-guttatus Rossi.
menavodi Vinson.
TI. List or SPECIES AND SUB-SPECIES RECOGNIZED IN THIS PAPER.
(For references and dates of synonyms, see Alphabetical List.)
1. Larropectus Hystrix E. Simon, 1889. (Plate XX VI. fig. 2.)
(Species. )
This form is quite distinct from any others that have been
taken in any part of the world, being clothed with short, stiff,
stout black spines, and cannot be mistaken for anything else.
Posterior row of eyes straight, approximately equidistant ; centrals
one diameter from each other, slightly further from the laterals.
Posterior centrals much larger than the anterior centrals. Central
anteriors less than one diameter apart, one diameter from the
laterals, and distinctly smaller. Laterals half a diameter apart.
The example figured was kindly lent by M. E. Simon, and
another example was found by Col. Yerbury at Aden.
Hab. Aden (Simon and Yertury).
9. LatropEctus cEomerricus C. L. Koch, 1841. (Plate XX VII.
ima, (fe) (Species. )
Synonym. L. zickzack (Karsch), sub Theridvum.
This form is more distinct from the others than any except
1 See note on p. 261.
1902. ] SPIDERS OF THE GENUS LATRODECTUS. 253
L. hystrix. See figures of the abdominal pattern. The central
anterior eyes are always, in all the examples which have come
before me, slightly larger than the laterals; and, though I am
unable to find any real difference in the form of the palpal organs _
of the male, yet the vulva of the female is distinctly different
from that of Z. mactans. The egg-cocoon is also characteristic,
being covered with small silky cusps, unlike that of Z. mactans,
which is of smooth silk.
This species | found commonly in the angles of windows in the
towns and villages throughout the Lower Amazons. Specimens
exhibit every variety of coloration from grey to black.
Hab. Sourn America: San Pedro and Rio Apia, Paraguay,
San Domingo and Curagao (Simon); Brazil (Keyserling); Rio
Janeiro (Goldi); Minas Geraes (Rogers); Lower Amazons, San-
tarem, ete. (7. P. Cambridge). Arrica: Khartoum (Voission) ;
Abyssinia, Mozambique, and Madagascar (Simon); Cape Colony,
Table Mountain (Hull); Jansenville (IMss Leppan); Cape Verde
Islands (Ff. P. Cambridge). Inp1a: Kurrachee and Manora
(Townsend). AustRauiA: Melbourne (ogg).
3. LATRODECTUS PALLIDUS O. P. Cambridge, 1872. (Plate XX VI.
fig. 1.) (Species. )
Hab. Plains of Jordan (0. P. Cambridge). Persian Gulf,
Bushire (Kurrachee Museum).
4, LavropEcTUS MACTANS (Fabr.), 1775. (Plate XX VII. fig. 2.)
(Species. )
Synonyms. Abbot’s Drawings: 191, 194, 195, 395, 344,—
L. formidabilis Walck.—L. variolus Walck.—L. intersector
Walck.—L. formidabilis Nicolet.—L. variegatus Nicolet.—
L. thoracicus Nicolet.—L. zorilla (Walck.), sub Tetragnatha.
—JL. dotatus C. L. Koch.—L. verecundum (Hentz).—L. line-
atum (Hentz).—ZL. apicalis Butler.—L. carolinus Butler.—
L. malmignathus, var. tropica Van Hasselt.
With regard to this form, after examining numerous examples
from North America and Central America and a few from Peru,
I have come to the conclusion that originally it was derived from
the same stock as tredecim-guitatus Rossi; and that whatever
small differences there are between the two now, they are the
result of long separation and different surroundings. The only
differences which appear to me to be constant, lie in the relative
position of the eyes of the anterior row and the hairy clothing of
the abdomen. It is true that the abdominal pattern is different,
the lateral spots being elongate-oval, or long narrow stripes, instead
of more rounded and shorter, as in tredecim-guttatus ; but since
there is every variety of coloration, from those which are entirely
black, or have only the apex of the abdomen red, to those which
are fully striped with red, one cannot regard colour-characters as
of specific importance.
254 MR. F. PICKARD CAMBRIDGE ON THE [Apr. 15,
5. LATRODECTUS CURAGAVIENSIS (Miiller), 1776. (Plate XX VII.
fig. 4.) (Sub-species. )
Synonym. L. geographicus, Van Hasselt.
The two females, referred by M. Simon to the former name,
from Asuncion, and kindly lent me for examination, are adult,
and represent a type of coloration often found in the immature
of Z. mactans. This form has the abdomen mainly red, with a
pair of parallel black longitudinal bands posteriorly, and two or
more transverse black cinctures anteriorly. The type of coloration
is the same as that of some males and females which I took on
the sandy campos near Santarem, on the Lower Amazons; and
the latter are unmistakably like Van Hasselt’s figure of L. geo-
graphicus, though the precise form of the coloration differs.
But, in this case again, though the Spiders are very much
smaller, Iam unable to find any really reliable difference in the
papal organs or in the vulva from those of Z. mactans. Differences
may indeed exist, and possibly with several dozen examples to
compare, instead of two or three, one might find them.
At present, however, I can only regard this form as a dwarf
race with different form of coloration, and for the time being
must consider it as a sub-species.
Hab. Curagoa, West Indies (Jfiller); Paraguay: Asuncion,
San Pedro and. Rio Apa (Simon); Surinam (Van Hasselt) ;
Amazons (J. Cambridge).
6. LarropEctuUs TREDECIM-GuTTATUS (Rossi), 1790. (Plate XX VI.
fig. 3.) (Sub-species. )
Synonyms. L. malmignaius Walck.—L. martius Aud. in Sav.
—L. argus Aud. in Sav.—JZL. .venator Aud. in Sav.—ZL.
erebus Walck.—L. lugubre (Dufour, sub . Theridion).—L.
oculatus Walck.—L. conglobatus C. L. Koch.—ZL. lugubris
Motsch.—L. hispida (C. L. Koch), sub Meta.
This is perhaps the best known form, being common all through
the South of Hurope, the Mediterranean region, Northern Africa,
and through South Russia and Syria. It is notorious in all parts
of this region for its venomous bite.
The only real distinction I can find. between the “ forma prin-
cipalis” of 13-guttatus and_mactans is that in the female the
lateral anterior eyes are further from the centrals than these are
from each other, and the hairy clothing is much finer. In the
males,. however, there is no difference in this respect. This.
character (the eye-formula), moreover, is not constant in the
females of either mactans, tredecim-guitatus, or hasseltii. I can
find no tangible difference between the various parts of the
palpal organs in the male, or of the vulva in the female, of these
forms (hasseltii_ 3 is unknown to me).
I believe these to be all offshoots of one original form, for which
the oldest name is macians. -
Hab. Kurore: Spain, Italy (Walck.); Greece (C. L. Koch) ;—
1902. ] SPIDERS OF THE GENUS LATRODECTUS. 255
France, Morbihan; Vendée; Vaucluse, Avignon; Hérault (Simon);
—Hungary, Croatia (Kulczynski) ;—Italy, Tuscany (Doria) ;—
Sardinia, Corsica (Walck., Simon) ;—S. Russia, Sarepta (Motsch.).
Asta Minor (Simon) ;—Arabia (Dr. Anderson) ;—Syria (Simon) ;
—Persian Gulf, Bushire (J/oore).
ATLANTIC IstANps: Canaries (Lucas) ;—Madeira, Porto Santo
(Kulczynski, Grant).
Arrica: Egypt, Alexandria (Walck., Aud. in Sav., Simon, etc.).
7. Larropecrus meNAvopI Vinson, 1863. (Plate XX VII. fig. 3.)
(Sub-species.)
This form, resembling the Australian examples (Aasseltii) in
size, but having the vestiges of the dorso-lateral spots character-
istic of 13-guttatus, consisting of small red spots, is peculiar to
Madagascar and the neighbouring islands. I am quite unable to
distinguish the female from either 13-guttatus or hasseltii, but so
far I have had no opportunity of examining the male. J am
inclined, however, to believe that it is simply a local variety of
13-guttatus, the Kuropean form of mactans.
Hab. Madagascar (Vinson).
8. LarropEctus HASSELTII Thorell. (Plate XX VI. fig. 4; Plate
XXVII. figs. 1, 6.) (Sub-species.)
Synonyms. ZL. scelio Thorell.—ZL. cinctus Blackwall.—L. indicus
Simon.—Z, elegans Thor.
This distinct-looking form is essentially Z. menavodi, which has
lost the small dorso-lateral spots, leaving only a narrower (Australia,
ete.), or broader and foliated (Loyalty Islands), red band down the
posterior dorsal central line.
Hab. Austratia: Rockhampton; Bowen (Port Denison); Cape
York (Thorell) ;—New Caledonia (Simon) ;—New Guinea (Z'urner);
—New Holland (Kulezynski) ;—Loyalty Islands, Lifu (Creagh) ;
—N. Britain (Brit. Mus.);—New 8. Wales, Hill Grove; N. Queens-
land, Muldiva (Broom); N.W. Australia (Beckett) ;—Adelaide
(Meldola).— Arrica: Shiré River; Zambesi (blackwall); Lake
Nyassa (Brit. Mus.); Bogos, Scioa or Shoa (Pavesi and Antinort)
Graham’s Town, Tea Fountain (a dark variety with only a minute
red apical spot and a few black blotches), Jansenville, Cape Colony
(Brit. Mus.). Asta: Persian Gulf, Bushire (J/oore); N.Guzerath ;
Poona; Mascat; Kurrachee (Simon and Brit. Mus.)— Burman
(fea).
9, Larropecrus KATIPO Powell. (Plate XX VII. fig. 5.)
(Sub-species. )
There is a decided tendency towards a shortness of the first
pair of legs amongst specimens from different parts ot Africa,
Australia, India, and New Zealand. This is most noticeable
amongst examples from New Zealand, and, together with the
tendency, the examples are as a rule smaller, and the anterior
256 MR. F. PICKARD CAMBRIDGE ON THE [ Apr. 15,
portion of the abdomen inclines to be marked by indistinct
narrow, transverse, pale cinctures, as in an example from Abyssinia ;
while the markings on those from New Zealand more resemble,
though very indistinctly, those of geometricus.
In the case of these examples there are unfortunately no males.
But although the first pair of legs in the New-Zealand examples
are distinctly shorter in proportion than those of L. hasseltti from
Australia, yet this character is found also, in varying degrees,
amongst examples from Australia and India, so that one cannot
regard it as constant.
Hab. New Zeauann: Portland Island, Hawkes Bay (obson) ;
Canterbury (Brit. Mus.).
III. Alphabetical List of Described or Figured Species of the
Genus LATRODECTUS.
Larropectus. Abbot’s figures 191 9, 194 9, 195 9, 395 ¢ ad.,
344 g juv. All these are varieties of LZ. mactans Fabr.
(Abbot’s drawings in Brit. Mus. Nat. Hist. 1792).
LATRODECTUS APICALIS A. G. Butler, 1877.
Proc. Zool. Soc. Lond. 1877, p. 75, pl. xiii. fig. 2, ete. Had.
Galapagos Islands. Type in coll. Brit. Mus.—A variety of
L. mactans Fabr.
LatropeEctus areus Aud. in Sav., 1825-27.
Savigny’s ‘ Egypte,’ p. 137, pl. 3. fig. 10. Hab. Alexandria.—
This is L. tredectm-guitatus (Rossi).
LatRoODECTUS CAROLINUS (A. G. Butler), 1877.
Proc. Zool. Soe. Lond. 1877, p. 75, pl. xui. figs, 3, 3a, 3 6.
Hab. Charles’s Island, Galapagos. Sub Vheridion.—An imma-
ture female of LZ. mactans Fabr.
LATRODECTUS cINCTUS Blackwall.
Ann. Mag. Nat. Hist. (3) xvi. p. 341 (1865). Hab. Shiré River,
South-east Africa, Zambesi. Also Bogos and Scioa (Shoa).—
This species is clearly a variety of the form called by Thorell
hasseltii. I have before me a young female from Shoa, referred
by Pavesi to “ cinctus,” which agrees with Blackwall’s description.
The form of the markings coincides also with that of Thorell’s co-
type of sceho from Cape York, which is now before me, the only
difference being that there is a single transverse bar beneath,
similar to that obtaining in the form called menavodi and also in
some examples of tredecim-guttatus. The shortness of leg i. in
proportion to the length of the carapace brings it nearer to the
katipo variety of hasseltit which is found in New Zealand, ete.
LATRODECTUS CONGLOBATUS C. Koch, 1838.
Die Arachniden, iv. p. 41, fig. 274. Hab. Greece.—A variety
of L. tredecim-guittaius Rossi.
1902.] - SPIDERS OF THE GENUS LATRODECTUS. 257
LATRODECTUS CURAGAVIENSIS (Miiller), 1776.
Linn, Vollst. Nat.-Syst. Supp., Reg. Band, p. 342. 1776 (sec.
Keyserling). Hab. Curagoa.—E. Simon, Boll. Mus. Zool. Anat.
Comp. Univ. Torino, vol. xii. no. 270, Feb. 4, 1897. Hab. As-
cuncion, San Pedro and Rio Apa, Paraguay.—This species, of
which M. Simon has kindly sent me examples, is, so far as color-
ation goes, similar to varieties of geographicus Hasselt. The
present name has priority.
LATRODECTUS CURASSAVICUM (Héring). Hab. Ozam.
(I am at present unable to indicate this reference.)
LATRODECTUS DisTINcTUS Blackwall, 1859.
Ann. Mag. Nat. Hist. (3) iv. p. 260. Hab. Madeira.—Belongs,
according to Thorell, to the genus Lithyphantes, possibly = L.
nobilis Thor.
Latroprectus botatus C. Koch, 1841.
Die Arachniden, viii. p. 115, fig. 683. Hab. North America.
Identical, according to Koch, with Tetragnatha zorilla Walck.
Both these forms are obviously males of LZ. mactans Fabr.
LATRODECTUS ELEGANS Thor., 1898.
Ann. Mus. Genova, (2) xix. p. 293, 9. Hab. Burmah.—
Probably a variety of Z. hasseltii; resembling L. cinctus Blackw.,
a form I have seen from Shoa.
LATRODECTUS EREBUS Walck., 1837.
Insectes Aptéres, i. p. 646. Hab. Egypt, Spain.—Dark variety
of L. tredecim-guttatus Rossi.
LATRODECTUS FORMIDABILIS Walck., 1837.
Insectes Aptéres, i. p. 647, 9. Hab. Georgia.— Undoubtedly
L. mactans Fabr.
LATRODECTUS GEOGRAPHICUS Van Hasselt, 1888.
Tijd. voor Ent. xxxi. pt. 3, pp. 165-200, pls. v. & vi. (1888).
Hab. Surinam.—This is, so far as size and pattern of colouring, a
distinct species. I have taken the adult male and female on the
sandy campo, at the roots of grass and under overhanging ledges,
near Santarem on the Lower Amazons. It is, however, difficult
to justify the apparent specific distinction by any tangible
structural difference. The form may finally be relegated to the
position of a local race, and perhaps be regarded as a sub-species.
Latropectus GEomETricus C, L. Koch, 1841.
Die Arachniden, viii. pp. 115-117, fig. 684.
This is perhaps one of the most distinct and easily recognizable
of the species, the larger size of the anterior central eyes being
constant. The colour varies from pinky grey to nearly black.
258 MR. F, PICKARD CAMBRIDGE ON THE [Apr. 15,
Latropectus HAsseLrit Thor., 1870.
Kongl. Svenska Akad. Forhandl, 1870, no. 4, p.369. L. Koch,
Arach. Austr. 1871, p. 276. Hab. New Holland; Rockhampton ;
Bowen (Port Denison).—This species is undoubtedly identical
with scelio Thor.
LAtTRODECTUS HIsPiDuS (C. Koch), 1836.
Die Arachniden, iii. p. 9, fig. 166. Sub Meta. Hab. Greece.—
Probably identical with L. tredecim-guttatus (Rossi). -
LaTRoDECTUS HYSTRIX E. Simon, 1889.
Ann. Soc. Ent. Fr. (6) x. p. 99 (1889). Hab. Yemen (Simon) ;
Aden (Yerbury).—A very distinct species, recognizable by its
straight posterior row of eyes, and abdomen clothed with short,
stiff bristles.
LaTRODECTUS INDICUS E. Simon, 1897.
Mém. Soc. Zool. Fr. x. p. 252 (1897). (scunio, var. indica.)
Hab. North Guzerath.—Bull. Mus. d’Hist. Nat. no. 7, p. 289
(1897). Hab. Mascat, Kurrachee.—Pocock, Arach. Ind. p. 23,
fig. 80 (1900). This is a variety of LZ. hasseliti = scelio Thor.
LATRODECTUS INTERSECTOR Walck., 1837.
Insectes Aptéres, i. p. 649. Hab. America.—Undoubtedly a
variety of Z. mactans Fabr.
Larropectus KATIPO Powell, 1870.
Trans. New Zealand Institute, tii. p.57. Hab. New Zealand.
The relatively shorter legs of the first pair, and the straightness
of the posterior row of eyes, in the female (I have not seen the
male), would suggest at first that this is a distinct species and not
identical with hasseliti Thor. = scelio Thor., but I find that this
character is not by any means constant. The hairy clothing on
the abdomen furnishes, however, a very good distinctive character.
LATRODECTUS LINEATUS (Hentz), 1850.
Boston Journ. Nat. Hist. vi. p. 281, pl. x. fig. 3.—‘ Spiders of
the United States,’ ed. Burgess, p. 153, pl. 17. fig. 3. Sub Theri-
dium. Hab. North America.—A variety of Z. mactans Fabr.
LATRODECTUS LUGUBRIS (Dufour), 1820.
Ann. Gén. Sei. Phys. iv. p. 355, pl. Ixix. fig.1. Sub Theridion.
—Identical with LZ. erebus Walck. & Aud. in Sav. = L. tredecim-
guitatus (Rossi).
LATRODECTUS MALMIGNATUS Walck., 1837.
Insectes Apteres, i. p. 642. Hab. Corsica, Sardinia, Italy.—
Variety of L. tredecim-guttatus (Rossi).
Latropecrus Martius Aud. in Sav., 1825-27.
Savigny’s ‘Egypte, p. 137. Hab. Italy (Walck.); Egypt—
1902.] _ SPIDERS OF THE GENUS LATRODECTUS. 259
Variety of L. tredecim-guttatus (Rossi)? According to Thorell
this belongs to the genus Lithyphantes.
LATRODECTUS MENAVoDI Vinson, 1863. '
Aranéides Réunion, ete. p. 122, pl. viii. fig. 5. Hab. Mada-
gascar.—This form may be regarded as a sub-species, or local
race.
LarropEctus ocuLAtus Walck., 1837. .
Insectes Aptéres, i. p. 645. Hab. Alexandria.—A variety of
L. tredecim-guttatus (Rossi).
LATRODECTUS ORNATUS Lucas, 1845.
Histoire d’Algérie, p. 233, pl. 14. fig. 8. Hab. Algeria.—-
According to Thorell it belongs to the genus Lithyphantes.
LaATRODECTUS PALLIDUS O. P. Cambr., 1872.
Proc. Zool. Soc. Lond. 1872, p. 287 (9). Hab. Plains of
Jordan (O. P. C.); Bushire, Persian Gulf (Kurrachee Musewm).—
The two females from the Kurrachee Museum are undoubtedly
identical with Z. pallidus O. P. C. The dull yellow colour of the
abdomen (which is almost entirely devoid of hairs), with the
slashed pattern on the sides and the conspicuousness of the four
impressed dots on the anterior dorsal area, renders it easily to be
recognized from any other form. The posterior row of eyes is
almost or quite straight, and the lateral anteriors are nearer the
centrals than these are to each other.
LATRODECTUS PERFIDUS Walck., 1837.
Insectes Aptéres, i. p. 647. Hab. Georgia.
tans.
Probably LZ. mac-
“LATRODECTUS QUINQUEGUTTATUS Krynicki, 1837.
Bull. Soc. Imp. Nat. Moscou, p. 75, tab. vi. fig. 2. Had. 8.
Russia.
Larropectus scevio Thorell, 1870.
Kongl. Svenska Vet.-Akad. Forhandl. 1870, no. 4, p. 370. L.
Koch, “Araneiden Australiens, 1871, p. 279. K. Simon, Mém.
Soc. Zool. Fr. x. p- 252, 1897. ‘El New Caledonia (Francois) :
Muscat, Kurrachee; and North Guzerath, Central Asia (var.
indica Sim.).—This form is undoubtedly identical with hasselti
Thor., the latter name taking priority.
Larropectus scuucui (C. L. Koch), 1836.
Die Avachniden, iii. p. 10, fig. 137. Hab. Greece. Sub Meta.—
Thorell, Kong]. Svenska Vet.-Akad. Hand. Band xiii. no. 5, p. 68.
E. Simon, Bull. Soc. Zool. Fr. ix. 1884, p. 21, note. Pavesi, Ann.
Mus. Genova, xv. p. 333 (1880), Tunis. E. Simon, Arach. France,
vol. v. p. 179 (1881), note.
I do not know this species. A male and two females, which
260 ON THE SPIDERS OF THE GENUS LATRODECTUS. [Apr. 15,
were kindly sent by M. Simon under this name, I am quite un-
able, at present, to separate from tredecim-guttatus. Simon thinks
(Avachn. France, vol. v.) that this form may be identical with
pallidus O. P. Cambridge.
Hab. Greece (C. L. Koch); Algeria, Spain, and Senegal (Simon) ;
Spain, Torrevieja (Zhorell); Shoa and Tunis (Pavesi).
LATRODECTUS SPINIPES Lucas, 1845.
Hist. d’Algérie, p. 233, pl. 14. fig.9. Hab. Algeria (Lucas).—
This is possibly either an Asagena or a Pecilochroa (Drasside).
LaTRODECTUS THORACICUS Nicolet, 1854.
Gay’s Histoire de Chile, p. 461. Hab. Chili (Wicolet).—Probably
a variety of Z. mactans (Fabr.).
LATRODECTUS TREDECIM-GUTTATUS (Rossi), 1790.
Fauna Ktrusca, vol. ii. p. 136, tab. ix. fig. 10.
This species is the common Mediterranean form of which ZL.
erebus Walck. is the dark unicolorous black variety.
Hab. Waly, Corsica, Sardinia, Spain, N. Africa, Greece, S. Russia,
Madeira, etc., etc.
LATRODECTUS TROPICUS Van Hasselt, 1860.
Tijdschrift voor Entomologie, 11. p. 46, pl. 5. Hab. Curagoa.—
Probably a small variety of L. geographicus Hasselt.
LATRODECTUS VARIEGATUS Nicolet, 1854.
Gay’s Histoire de Chile, p. 461, pl. 4. fig. 9. Hab. Chili.—
Probably a variety of LZ. mactans Fabr.
LatropEctus vARIOoLUS Walck., 1837.
Insectes Aptéres, i. p. 648. Hab. Georgia.—Variety of L. mac-
tans Fabr.
LaTRODECTUS VENATOR Aud. in Sav., 1825-27.
Savigny’s ‘Egypte,’ p. 138, pl. 3. fig.11. Mab. Egypt.— Variety
of L. tredecim-guttatus (Rossi).
LATRODECTUS VERECUNDUS (Hentz), 1850.
Boston Journ. Nat. Hist. vi. p. 280, pl. x. figs. 1 & 2. Sub
Theridion. Hab. N. America.—A dark variety of LZ. mactans
(Fabr.), having the same relation to it as LZ. erebus bears to
L. 13-quitatus.
LATRODECTUS ZICKZACK (Karsch), 1878.
Zeits. gesammt. Naturwiss. li. p. 311 (1878). Sub Theridiwm.
Hab. Zanzibar.—Variety of L. geometricus C. Koch.
LaTRODECTUS ZORILLUS (Walck.), 1837.
Insectes Aptéres, Atlas, pl. 19. figs. 28, 2d, 2D. Sub Tetra-
gnathe zorille, § ad.—Obviously L. mactans Fabr.
1902.] ON THE PAINTED SNIPE AND PHEASANT-TAILED JAGANS, 261
IV. EXPLANATION OF THE PLATES.
Pratt XXVI.
Fig.1. Latrodectus pallidus; Bushire: p. 253. Dorsal view of female. la. A
portion of the integument from the lateral region of the abdomen, showing
acanthoid spines.
2. Latrodectus hystrix; Yemen: p. 252. Dorsal view of female. 2a. Integu-
ment of abdomen, showing acanthoid and stouter straight spines.
3. Latrodectus tredecim-guttatus; Tuscany: p. 254. Dorsal view of female.
3a. Integument of abdomen, showing acanthoid and long curved spines.
4, Latrodectus hasseltii: p.255 (co-type of L. scelio Thor., Cape York). Dorsal
view of female. 4a. Integument of abdomen, showing acanthoid and
longer spines. 46. Dorsal view of female variety from New Britain. 4c.
Another variety from the same locality. 4d. Another variety from Fort
Bowen. 4e. Variety from South Australia. 4 Variety (probably L.
cinctus Blackw.) from Shoa, Africa. 4g. Variety from Lake Albert Nyanza,
Africa,
Pratt XXVII.
Fig.1. Latrodectus hasseltii, var. indicus; Kurrachee: p. 255. Dorsal view of
female. la. Integument of central dorsal posterior red band, showing
absence of acanthoid spines. 16, 1c,1d,1e. Varieties from the Loyalty
Islands. 1. Integument of central dorsal posterior band, showing bristles
set closer together than in var. indicus.
2. Latrodectus mactans; Mexico: p.253. Dorsal view of female. 2a. Integu-
ment of abdomen, showing fine bristles only.
3. Latrodectus menavodi; Madagascar: p. 255. Dorsal view of female. 3a.
Integument of abdomen, showing fine bristles of two lengths.
4, Latrodectus curacaviensis; Santarem, Lower Amazons: p.254. Dorsal view
of female from Santarem. 4a. Variety from Curagoa, almost identical in
coloration with other varieties from Santarem.
5. Latrodectus katipo; New Zealand: p. 255. Dorsal view of female. 5a.
Ditto, variety. 56. Ditto, variety. 5c. Ditto, variety. 5d. Profile view,
showing relative length of leg i. 5e. Integument of abdomen, showing fine
hairs only.
6. Latrodectus hasseltii: p.255. Profile view, showing relative length of leg i.
7. Latrodectus geometricus; Cape Town: p. 252. 7a, 7b, 7c, 7d, 7¢. Dorsal
view of females, varying in colour from almost white to almost entirely
black, from Table Mountain.
3. Notes on the Painted Snipe (Rostratula capensis) and
Pheasant - tailed Jacana (flydrophasianus chirurgus).
By Frank Finy, B.A., F.Z.S., Deputy Superintendent
of the Indian Museum, Calcutta.
[Received March 12, 1902. ]
Few Waders are better known to sportsmen and to other
observers of wild bird-life in India than these two beautiful
species. But there is always something to be learnt from the
close observation necessary to the successful retention of
specimens in captivity; and as I have made a special study of
examples of these two birds with a view to sending them to the
Society’s Gardens—in which I was fortunately successful ’—I
venture to record a few notes which I have been led to make
concerning them.
1 [Three Painted Snipes were received from Mr. Finn on January Ist, 1901, and
nine Pheasant-tailed Jacands on January 11th, 1902 (see P. Z.S, 1902, vol. i. p. 61).
—Ep. |
yA) MR. F, FINN ON THE PAINTED SNIPE [ Apr. 15,
I. The Painted Snipe.
This species, although not so abundantly brought into the
Calcutta Provision Market as the true Snipes Gallinago celestis
and G'. stenura, is nevertheless commonly to be had during the
winter, and I have kept many examples. They bear confinement
well on the whole, but never become properly tame, although
they will let themselves be caught more readily than most birds.
This is no doubt largely due to the fact that they are in the habit
of either crouching to avoid detection—their olive-green dorsal
coloration, with yellow longitudinal stripes, making for protection—
or of endeavouring to frighten off an otherwise unavoidable enemy
by hissing and expanding their spotted wings. When only
slightly alarmed they open the wings without spreading them ; if
still more persecuted, they expand the further wing and raise
it ; while, in desperation, they spread both wings and the tail,
forming a most beautiful fan. Both sexes do this equally, and,
so far as I can judge, make much the same sound, which is like
that produced by plunging a hot iron into water. At the same
time they crouch down close to the ground, I have seen them
show off in this way to other. birds—a Rail, a Ruff, and a Pitta,
none of which were at all impressed by the display. I have,
however, seen a Golden Plover (Charadrius fulvus) frightened
thereby, although this species is bold and pugnacious with other
birds.
T have no doubt that the natives who informed Hume that
this was also the position assumed during courtship were correct
in their statements, since in so many birds the so-called courting
postures are merely those assumed on any excitement, as may be
well seen in the Turkey and Muscovy Duck.
When at rest the Painted Snipe squats down with the breast
on the ground and the tail up, the bill also pointed downwards.
At such times the head has a smooth rounded appearance. When,
however, the bird is moving about, there appear two superciliary
ridges, which give the head a quite different expression and
show off the magnificent dark eyes. There is also during move-
ment an up-and-down motion of the hinder part of the body,
similar to that observable in the Common Sandpiper (7Zringoides
hypoleucus), but slower.
This so-called Snipe appears never to bore for food, but it will
search for it in water, or even sand, with a motion much like
that of the Spoonbill. It eats grain—paddy and canary-seed—
readily, the latter seeming to be best assimilated, although the
former is more readily taken. It will also devour maggots, but
appears not to care about worms. The flight is entirely Rail-like,
and when the bird is skulking along to hide, the gait is also like
that of a Rail. Like a Rail, also, the Painted Snipe swims
readily, but this of course is not an important point, as the power
of swimming is general among the Waders. The grain-eating
habit is also common to the Ruff and Black-tailed Godwit among
the family allies of Hostratula.
1902.) AND PHEASANT-TAILED JAGANA, 263
I once kept a young bird of this species, which I obtained
when nearly full-fledged. 1+ fed well on canary-seed and maggots,
and I kept it till it was full-grown, but it showed no more
tendency to become tame than an old bird.
I am rather astonished at Blyth’s failure to keep this species
alive, as I do not consider it hard to manage, except for its
tendency to fly against netting and to abrade its bill by its futile
attempts to escape. If kept in a cage with upright bars it is
certain to hurt itself, but in a hutch with netted front, or a good-
sized aviary, it will do well, and some have survived for a year at
the Calcutta Zoological Gardens, though several succumbed after
this on getting access to unsoaked grain.
Il. The Pheasant-tailed Jacana.
This very beautiful and graceful wader has been a special
favourite of mine ever since I began seriously to study it. It is
one of the most numerously-captured species during the winter,
but it is not by any means easy to keep in captivity. The
difficulty lies in the fact that the birds’ legs and feet must be
kept damp in order that they may thrive, as otherwise the skin
about their hocks cracks and dries, and they become lame.
I used to turn out the birds that did not seem to be doing well
on the pond in the Museum grounds, generally with clipped
wings, and several lived there for some time, remaining most of
the time on masses of ‘“kalmi” or water-convolvulus.. Three
males are still (February) there, two of which are full-winged, but
seem to have no desire as yet to go away. When standing on
the weeds, they are most difficult to see from the other side of the
pond, which is about sixty yards across, but on the wing they are
most conspicuous objects. In this respect they much resemble
the Paddy-bird or Pond-Egret (Ardeola gray), which has a similar
plan of coloration, with a brown upper surface and concealed
white wings. Both birds have a somewhat similar flight, and,
were they insects, one would probably be said to mimic the other.
No doubt, however, in both birds the coloration is merely pro-
tective. When in breeding-plumage both the Egret and the
Jaganad are easy to see, especially the latter, which is a very
conspicuous object in the pied livery of the nuptial season, set off
by the long black tail. A female on the tank last May had
assumed nuptial plumage, as had one male, while two other males
and a pair of young birds still remained in undress.
The old hen would only allow one male to remain near her,
this being one of the winter-plumaged adults ; the full-plumaged
male was not allowed to approach, and even the favoured bird,
although he often drew near her in a stooping attitude, which
was the nearest approach to courtship I saw, was driven off if he
got toonear. The birds never made any display of their beautiful
white black-bordered wings, which rather surprised me. I found
that the birds were distinctly pugnacious about this time, observing
264 ON THE PAINTED SNIPE AND PHEASANT-TAILED JAGANA. [ Apr. 15,
boundaries strictly and attacking strangers. In fighting, they
seize with the beak and strike with both wings at once. The spur
with which the carpal joint is armed is only represented by a
small, movable, pointed tubercle in winter, and is evidently a
seasonal growth like the horns of the deer. They swim gracefully,
but slowly. Of many birds turned on the pond, only two were
seen to dive in trying to get away when first let out. They swam
only with their wings, which are remarkably powerful, very
unlike those of the Rails. The feet are very weak in grasping-
power, the hallux especially only flexing at the basal joint ; in the
Rails the last joint flexes strongly.
There is a good deal of variation in this species. I have seen
one or two specimens with particularly stout strong bills, fine old
females; some, irrespectively of age or sex, are glossed with
purple on the brown upper surface at all times—all show the
gloss when wetted. The eyes are usually brown in adults and
yellow in the young; but I have seen two quite young birds with
the dark iris, and many old ones, in adult winter plumage,
retaining the yellow eye. Birds showing the iris in a state of
change are curiously rare.
In watching the market for years, I never saw an adult retaining
its summer plumage in winter; but oneof my males on the pond,
which has “stuck in the moult,” to use a bird-fancier’s expression,
has never thoroughly changed into winter dress. From obser-
vation of another of these males, I believe the quills are cast
at once.
This bird does not appear to fear Kites, nor do these offer to
attack it, even when sickly; perhaps they fear its strong and
armed wings. It is a bold species, caring little for other birds,
and not timid with man after a few days’ immunity on a pond;
in a cage it is not so easily reconciled, and is apt to hurt its wings
at the carpal pot, much as Doves will frequently do when first
caged. Almost the only food I have seen taken by the birds at
liberty is small water-snails about the size of peas; in captivity
they will take to paddy at once, and also eat canary-seed and
maggots. The grain should always be soaked for them, and their
legs frequently wetted when they are confined in a cage. They
never seem to feel the heat in the open, remaining in the hot sun
all day long without panting.
The males are certainly better protected than the females, the
greater height of which exposes their white breasts when on the
low kalmi weed; but no doubt in more lush vegetation the hens
also would share in the protection which the coloration affords to
this most interesting bird.
PZ S.NS02 well Pl ZOO.
PJ.Simit delet hth. : thntern Bros.imp.
12 MICRALE STES STORMSI. 3 PHRACTURA LINDICA.
M3
ae
bs ceitiia
Ba
30 EES
ree
IP, 8) IVO2 areal Pl OUK. .
Mantern Bros .imp-
1. AUCHENOGLANIS PUNCTATUS.2.AUCHENOGLANIS PULCHER.
PB.J.Smit delet hth
S JAMME SOUL INS: BRE WLS
TOW WORE WEELILIS Wo) MLS Shel
USMS Wiley Wh SG “Sava VIN ViNOS SelOwS EA vaGGlats al" I
‘dum’ sorg usequry MIRME FO ENO SSessy Ie ist
DOO Nel
1902.] ON THE ICHTHYOLOGY OF THE CONGO. 265
\
4. Contributions to the Ichthyology of the Congo.—II. On
a Collection of Fishes from the Lindi River. By G. A.
Bovuencer, F.R.S8.
[Received March 17, 1902. ]
(Plates XX VIII.-XXX.")
Among the material which has been most obligingly intrusted
to me for study by the Director of the Royal Natural History
Museum in Brussels there is a large and important collection of
Fishes made by M. Maurice Storms, a cousin of the late Raymond
Storms, so well known for his important contributions to pals-
ichthyology, in the Lindi River, which flows into the Congo at
the Stanley Falls. As the fauna of this river had not previously
been explored, it is desirable to give a list of all the species repre-
sented in the collection, seven of which are new to science.
Mormynrip&.
Mormyrops DELICIosus Leach.
. PETROCEPHALUS SIMUS Sauv.
. MARrcUSENIUS PULVERULENTUS Blgr.
SroMATORHINUS HUMILIOR Bler.
Myomynrus MACRODON Bier.
GNATHONEMUS Moor Gthr.
. GNATHONEMUS ELEPHAS Bler.
cost Ot go tO
GNATHONEMUS BHYNCHOPHORUS Bler.
CHARACINID A.
9. Hyprocyon Lineatus Blkr.
10. BrycoN#THIops microstomMAa Gthr., var. BOULENGERI,
Pellegr.
11. ALESTES GRANDISQUAMIS Bler.
12. MicRALESTES HUMILIS Bler.
13. Micraestes autus Bler.
14. MIcRALESTEs sToRMSI, sp.n. (Plate XXVIII. figs. 1 & 2.)
Depth of body 34 times in total length, length of head 4 times,
Head longer than deep, twice as long as broad; snout shorter
than the eye, the diameter of which equals the interorbital width
1 For explanation of the Plates, see p. 271.
Proc. Zoot. Soc.—1902, Vou. I. No XVIII. 18
266 MR. G. A, BOULENGER ON THE [ Apr. 15,
and is contained 23 to 3 times in the length of the head;
maxillary extending “to below anterior border of eye; premaxillary
teeth 16, in two rows, the outer tricuspid, the inner quinque-
cuspid ; mandibular teeth 8 in the outer row, tricuspid. Gull-
rakers short, 12 or 13 on lower part of anterior arch, Dorsal IT 8,
originating above ventrals, at equal distance from the tip of the
Spout and the root of the caudal, 13 as deep as long and about
3 the length of the head. Adipose fin 2 to 3 times as distant
from the rayed dorsal as from the caudal. Anal III 15-16,
deeper and with more convex border in the males than in the
females. Pectoral longer than the ventral, # or 4 the length of
the head. Caudal forked, with peor lobes. Caudal peduncle
as long as deep. Scales 22-24 ap 2 between lateral line and base
of ventral. Brownish above, white beneath, with a silvery
lateral band,
Total length 75 millim.,
Numerous specimens.
15. DisticHopus FASCIOLATUS Bler.
16. DisTICHODUS SEXFASCIATUS Blegr.
17, NANNOCHARAX FASCIATUS Gthr.
18. NanNocHARAX ELONGATUS Blgr.
19. NANNOCHARAX TENIA Bler.
CYPRINIDE.
20. LaBro GREENII Blegr.
21. Laseo parvoes Bler.
22. BARBUS KESSLERI Stdr.
23. BARBUS HUMERALIS Bler.
24. BARILIUS UBANGENSIS Pellegr.
B. fasciolatus Blgr. is identical with this species. The fascicle
of the Bulletin du Muséum containing Dr. Pellegrin’s description
was not received in London until March 3rd of this year, after
my description had gone to press in the Ann. Mus. Congo, Zool.
ii. p. 34 (March 1902).
25. CHELETHIOPS ELONGATUS Blgr.
SILURIDEZ.
26. CLARIAS ANGOLENSIS Stdr.
27. CLARIAS BYTHIPOGON Sauv.
28. EurTRoPIuS CONGOLENSIS Leach.
1902. ] ICHTHYOLOGY OF THE CONGO, 267
29, AUCHENOGLANIS PUNCTATUS, Sp.n. (Plate X XIX. figs. 1, 1a.)
Depth of body 5 times in total length, length of head 3 times.
Head smooth, 14+ as long as broad; snout pointed, half the length
of the head; diameter of eye 6 times in the length of the head,
hardly twice in the interocular width; width of mouth more
than half that of the head ; lips wide, papillose; posterior nostril
cleft-like, a little nearer the eye than the end of the snout; pre-
maxillary teeth forming a short and narrow band, the mandibulars
two rounded groups; maxillary barbel as long as the head,
reaching the middle of the pectoral spine; outer mandibular
barbel 14 the length of the head, inner 4 the length of the head ;
gill-membranes forming an obtuse angle ; occipital process small,
as long as the eye, separated from the interneural shield, which
is very small. Humeral process small, pointed. Dorsal I 7;
spine smooth, 2 the length of the head, Adipose fin 4 times as
long as deep, nearly reaching the caudal, narrowly separated from
the rayed dorsal. Pectoral spine half as long as the head, its
inner edge strongly serrated. Ventral not reaching the anal.
Latter with 10 rays, 7 of which are branched. Caudal rounded.
Pale brownish, with seven indistinct darker bars, each accom-
panied by a vertical series of black dots; belly white; dorsal and
candal fins with transverse series of dark spots.
Total length 80 millim.
A single specimen.
Allied to A. ubangensis Blgr. Differing principally in the
longer maxillary barbel.
30, AUCHENOGLANIS PULCHER, sp.n. (Plate X XIX. figs. 2, 2 a.)
Depth of body 5 to 5} times in total length, length of head
3 to 35 times. Head smooth, 13 as long as broad ; snout obturely
pointed, half the length of the head; diameter of eye 7 to 9 times
in length of head, twice to twice anda half in the interorbital
width ; width of mouth rather more than half that of the head;
lips wide, papillose; posterior nostril cleft-like, a little nearer
the eye than the end of the snout; premaxillary teeth forming a
reniform group, the mandibulars two rounded groups; maxillary
barbel $ to # the length of the head, not extending beyond the
base of the pectoral ; outer mandibular barbel nearly as long as the
head, inner nearly half the length of the head; gill-membranes
forming an acute angle; occipital process small, as long as the
eye, separated from the interneural shield, which is very small.
Humeral process small, pointed. Dorsal 17; spine smooth, half
the length of the head. Adipose fin 4 to 5 times as long as deep,
twice to twice and a half as long as its distance from the rayed
dorsal. Pectoral spine half as long as the head, its inner edge
strongly serrated. Ventral not reaching the anal. Latter with
11 or 12 rays, 7 or 8 of which are branched. Caudal rounded.
Caudal peduncle as long as deep. Yellowish, brown on the back,
with transverse series of round black spots; a large black blotch
18*
268 MR. G. A, BOULENGER ON THE (Apr. 15,
on each side below the dorsal fin; belly white; dorsal and caudal
fins with round black spots.
Total length 100 millim.
Several specimens.
Distinguished from the preceding species and from A. uban-
gensis by the smaller eye ; from the former by the shorter, from
the latter by the longer maxillary barbel.
31. AMPHILIUS’ BREVIS, sp.n. (Plate X XIX. fig. 3.)
Depth of body 6 to 64 times in total length, length of head
35 to 32 times. Head much depressed, a little longer than broad ;
snout broadly rounded, 3 the length of the head ; eye very small,
9 or 10 times in feng of head, 23 or 3 times in interocular
width; maxillary barbel $ or ? length of head, scarcely longer
than the outer mandibular, meer mandibular about 3 length of
head. Dorsal I 6, equally distant from the end of the snout and
the root of the caudal ; adipose fin once and a half to twice as
long as the rayed dorsal, 13 to 14 as long as its distance from the
latter. Pectoral longer fiom he ventral, ¥ the length of the
head. Anal JI 6. Caudal feebly emarginate. Caudal peduncle
as long as deep. Brown above, dotted with black; belly white ;
caudal peduncle blackish towards the base of the caudal; fins
white, caudal with a large rhomboidal or cruciform black marking.
Total length 48 millim.
Two specimens.
This species is most nearly related to A. platychir Gthr., which
differs, among other points, in having the dorsal fin nearer the
end of the snout than the caudal, and the caudal peduncle longer
than deep.
32. SYNODONTIS GRESHOFFI Schilth.
33, SYNODONTIS PLEUROPS Blgr.
34, SYNODONTIS DECORUS Blgr.
35, HUCHILICHTHYS ROYAUXI Blgr.
36. PHRACTURA LINDICA, sp.n. (Plate XXVIII. figs. 3, 3a, 3.)
Depth of body 74 to 8 times in total length, length of head
5 to 53 times. Head 13 to 12 as long as broad, nearly smooth
above, covered with papillose skin; snout half length of head,
obtusely pointed, projecting but slightly beyond the mouth; space
between the two nostrils at equal distance from the end of the
snout and from the eye, or a little nearer the latter; eye supero-
lateral, its diameter 6 to 7 times in length of head, twice in inter-
ocular width; barbels thick and papillose, annulate, maxillary
z length of head, outer mandibular 3, inner mandibular 3; occi-
pital process narrow, 4 times as longas broad, narrowly separated
1 Amphilius Gthy. 1864, = Anoplopterus Pfeff.
1902. ] ICHTHYOLOGY OF THE CONGO, 269
from the small interneural shield. Dorsal I 6, first ray longest,
slightly longer than the head ; second dorsal very small, originating
above last rays of anal, its posterior rays adnate to the back
through a transparent membrane. Anal II 7-8. Pectoral a
little longer than head, reaching, or not quite reaching base of
ventral; latter a little shorter, reaching anal; outer ray of
pectoral and ventral much thickened. Caudal with crescentic
notch. Caudal peduncle much depressed, # to 3 total length.
23 or 24 dorsal and 18 or 19 ventral scales, of which 8 or 9 are
on the caudal peduncle, the last 5 united round the latter.
Yellowish brown above, speckled with darker and with four more
or less distinct broad dark cross-bands; fins whitish, with brown
spots forming bars across the pectorals and ventrals.
Total length 82 millim.
Four specimens.
The genus Phractura Blgr. was represented by three species :
P. bovet Perugia, from the Lower Congo, of which I have lately
examined three specimens belonging to the Brussels Museum ;
P. scaphirhynchura Vaill., known from two specimens from the
Alima, Upper Congo; and P. ansorgit Blgr., recently described
in these ‘ Proceedings’ from a single example discovered by
Dr. Ansorge in Southern Nigeria. The four species now known
may be distinguished by means of the following synopsis :—
I. Occipital process not reaching interneural shield ;
snout about half length of head.
Head feebly rugose above; posterior nostril nearly
as distant from the eye as the anterior from
the end of the snout; diameter of eye 10 or
11 times in length of head, 2} or 3 times in
interocular width; maxillary barbel $ length
OF MEAGRE eseagncee oan oate (eebeee horsemen, Lev. Gover Poragia:
Head smooth above; posterior nostril nearly as
distant from the eye as the anterior from the
end of the snout; diameter of eye 6 or 7
times in length of head, twice in interocular
width; maxillary barbel 4 length of head ... 2. P. lindica Blgv.
Head rugose above, with strong ridges ; posterior
nostril very near the eye, the diameter of
which is 7 times in Jength of head and 13 in
interocular width; maxillary barbel $ length
GE INVENtl ccoedocosnbe donocahspacsnccdccosascncocbecdtece: ten Jan Qornead Lei laie
IL. Occipital process in contact with interneural
shield; snout more than half length of head ;
eye 7 times in length of head, 13 in interocular
width ; maxillary barbel 4 length of head ...... 4. P. scaphirhynchura Vaill.
CYPRINODONTID.
37. HAPLOCHILUS sINGA Blgr.
ANABANTID A.
38. ANABAS MACULATUS Thomin.
270 MR. G. A, BOULENGER ON THE [Apr. 15,
CICHLIDA.
39. HEMICHROMIS FASCIATUS Peters.
40, PSEUDOPLESIOPS SQUAMICEPS, sp.n. (Plate XXX. fig. 1.)
Depth of body 33 times in total length, length of head 3 times.
Upper profile of head curved; snout a little longer than the
diameter of the eye, which is contained 33 times in the length of
the head and equals the interorbital width; mouth with broad,
thick lips, extending to below anterior border of eye; 3 series of
teeth in the upper jaw; occiput and sides of head with large
scales; 2 or 3 series of scales on the cheek. 8 gill-rakers on lower
part of anterior arch. Dorsal XVII 8; spines subequal from the
sixth, barely one third length of head; longest soft rays two
thirds length of head, Anal III 6; third spine longer than
dorsals, two fifths length of head; soft rays like dorsals. Pectoral
about two thirds the length of the head. Ventral about the same
length, not reaching the vent. Caudal rounded. Caudal peduncle
as long as deep. Scales cycloid, 29 in a longitudinal, 13 or 14 in
a transverse series; upper lateral line on 8 or 9 scales, forming
an interrupted series, lower on 3 or 4, Pale brownish above,
white beneath ; a dark horizontal streak on each side of the head,
behind the eye; soft dorsal, anal, and caudal fins with numerous
transverse series of small dark spots.
Total length 65 millim.
Two specimens.
Distinguished from P: nudiceps by the scales on the head, the
shorter dorsal spines, and the interrupted lateral lines.
4], TILAPIA sTORMSI, sp.n. (Plate XXX. figs. 2, 2a.)
Teeth in 4 to 7 series, outer deeply notched, inner much smaller
and tricuspid; 36 to 60 teeth in the outer premaxillary series.
Depth of body equal to length of head, 3 to 33 times in total
length. Snout rather pointed, with straight or slightly convex
upper profile, 13 to 12 the diameter of the eye, which is contained
4 to 43 times in the length of the head and equals the inter-
orbital width; width of mouth about 2 that of the head;
maxillary extending to between nostril and eye; 3 or 4 series of
scales on the cheek; large scales on the opercle. Gill-rakers
short, some anvil-shaped, 11 or 12 on lower part of anterior arch.
Dorsal XVI-XVIT 9; last spine longest, about 2 the length of
the head, about # the longest soft rays. Pectoral rounded, about
% the length of the head, widely separated from the anal.
Ventral not reaching the vent. Anal II] 7; third spine about
1 the length of the head. Caudal rounded. Caudal peduncle
as long as deep. Scales with strong marginal denticulation,
22-3 a qe aes :
30-31 [7473 lateral line Sir scales on the occiput and nape very
small. Olive-brown above, some of the scales black at the base;
5 or 6 more or less indistinct dark cross-bands; a blackish
1902.] ICHTHYOLOGY OF THE CONGO. _ 271
opercular spot; dorsal and caudal fins with small dark spots ;
a fine blackish edge to the caudal above and beneath.
Total length 102 millim.
Five specimens.
Allied to 7’. fasciata Perugia.
Appendix.
In a series of specimens from other parts of the Congo State,
belonging also to the Brussels Museum, the representative
of a new species of a marine Clupeid genus was found, which I
have great pleasure in naming after my friend and colleague
M. L. Dollo.
PRISTIGASTER DOLLOI, sp. n. (Plate XXX. fig. 3.)
Closely allied to P. cayanus Cuv., but abdominal profile much
less convex. Depth of body 3 times in total length, length of
head 4 times. Hye longer than the snout, shorter than the post-
ocular part of the head; maxillary extending to below the centre
of the eye. Gill-rakers long, 25 on lower part of anterior arch.
Dorsal 15, equally distant from the end of the snout and the root
of the caudal. Anal 46, originating below the middle of the
dorsal. Pectoral nearly as long as the head. Caudal deeply
forked. Caudal peduncle as long as deep. About 40 scales in a
longitudinal series. Ventral serration formed of 32 spinose scutes.
Uniform silvery.
Total length 120 millim.
A single specimen from Banana.
EXPLANATION OF THE PLATES.
Prats XXVIII.
Fig. 1. Micralestes stormsi, male, p. 265.
2.
Bs female.
3. Phractura lindica, p. 268.
3a. sb “ Upper view of head and anterior part of body, X 2.
35. c on Lower view of head, X 3.
Puate XXIX.
Fig. 1. ct oe punctatus, p. 267.
la Hs Open mouth, X 2.
2. cf es p. 267.
2a. a Open mouth, X 2.
3. Amphilius brevis, p- 268.
Puate XXX.
Fig. 1. Pseudoplesiops squamiceps, p. 270.
2. i eetae i? stormsi, p. 270.
Open mouth, X 2.
3 pera cgaster ’ dolloi, p- 271.
272 MR. H. PRICHARD ON PATAGONIAN MAMMALS. [ Apr. 15,
5. Field-Notes upon some of the larger Mammals of Pata-
gonia, made between September 1900 and June 1901.
By Hesxera Priowarp, F.Z.S.
{Received March 18, 1902. ]
1, Tue Huemun. (YXenelaphus bisuleus.)
(Huemul of the Argentines and Chilians ; Ciervo of the Gauchos
of Southern Patagonia; Shoan of the Tehuelches.)
In the neighbourhood of Lake Buenos Aires this beautiful
deer first came under my observation. On the south side of the
valley of the river De los Antiguos I saw a buck (which I shot),
two does anda pricket. I wasinformed by my Gaucho, Humphrey
Jones, that the Huemul is found in the woods as far north as the
Welsh Colony of the 16th October about lat. 43°, and that on
the south its range extends to the Straits of Magellan. Its
present habitat may be broadly said to extend as far east as the
foothills of the Andes. Dr. F. P. Moreno states that the Xene-
laphus bisulcus has been seen in the hills in the vicinity of Port
Desire on the Atlantic coast; I do not, however, think it is any
longer to be found there. So far as my personal observations go,
I never came across a specimen farther east than a couple of
miles from the shores of Lake Buenos Aires upon its north-
eastern side. The Indians say that this animal was at one time
more numerous in this region.
During the summer these deer leave the lower grounds where
the mosquitoes trouble them, and travel up to the snow-line of
the Cordillera and even beyond it. At this season I never
observed a large herd; but in the winter, Mr. Cattle, a pioneer
living near Lake Argentino, informed me that a numerous herd,
over 100 strong, had visited the lake.
The Huemuls are in the habit of wandering outside the forests
in the morning and forenoon, but in the afternoon they generally
retire to their shelter, where they often lie down. I have found
these animals in the dense forests upon the slopes of the Cordillera
which border the lakes. They are excellent swimmers, and cross
the broad arms of Lake Argentino without hesitation.
In December the Huemuls which I shot were shedding their
winter coat, and I noticed that the bucks were further advanced
in this matter than the does. There were a few scraps of velvet
clinging to the horns of one of the bucks which I shot on the
17th of December.
The best head that I secured carried 5 points. Mr. Von Plaaten
Hallermund, of the Argentine Boundary Commission, told me
that he had seen a Huemul’s head carrying 8 points; this was in
the neighbourhood of Lake San Martin. One of my peones,
Bernardo Hahansen, who had penetrated into the same district,
said he also had seen an 8-pointer. Mr. Cattle and his companions
shot two bucks, both of which were 4-pointers.
1902.] MR. H. PRICHARD ON PATAGONIAN MAMMALS. 273
Save for the attacks of the Pumas, Xenelaphus bisulcus lives
pretty well undisturbed in its fastnesses. The Indians do not
hunt the Huemul, as in the forest-land their horses and boleadores
are comparatively useless. They do occasionally kill a few, which
may have strayed to the foothills and to the shores of the lakes.
These deer, which know little of man, are in general very
confiding. Near the Colony of the 16th October, Jones told me
that they had become very wary and difficult of access, as was to
be expected in a region where they are constantly hunted. In
the unpenetrated districts the buck is very courageous in the
rutting-season, and has been known to make some show of
attacking man. On open ground, according to my experience,
they showed wonderfully little timidity, and would wait the
approach of man, but inside the forests they invariably dashed
away on catching a glimpse of one of our party.
“When it has observed something unusual in its surroundings,
the Huemul will remain watching and without moving for a great
space of time. On one occasion, I saw near Lake Argentino a
buck and doe about a quarter of a mileaway. I lay undera bush,
watching some wild cattle, and the Huemuls stood and watched
me for nearly an hour. They were about 10 yards apart. On
my returning to the same spot in the evening, I found them still
watching my horse, which I had tied up in their view.
In one or two instances, when I fired at a Huemul, the others
of the herd have run towards the noise. Once this occurred
when I was in full sight of the animals. If, however, you have
a dog with you, they will immediately take to flight.
Musters, in his ‘ Travels in Patagonia,’ mentions a ‘‘red” deer.
Of this I could find no trace; so that I conclude that he probably
referred to Xenelaphus bisulcws under this name.
2. Toe Puma. (felis concolor puma.)
(Zion of the English settlers; Leon of the Argentines, Chilians,
and Gauchos ; Grol of the Tehuelches.)
The distribution of Felis concolor puma extends over the entire
country of Patagonia. It is to be found in the Cordillera as on
the pampas. I came upon tracks of this animal at the end of the
north-west arm of Lake Argentino, about long. 73° 14’, and I
also saw a Puma at the south-western extremity of that lake.
Evidence of their existence accompanied the whole itinerary of
the expedition throughout the entire route it covered. The
number of Pumas in Patagonia is very great, more so than any
traveller has as yet given any idea of. Two pioneers killed 73
in one winter near Lake Argentino. Near San Julian immense
numbers are yearly destroyed, but now, owing to the advent of
settlers, their numbers are decreasing. At Bahia Camerones, on
the farm of Mr. Greenshields, 14 Pumas were killed during the
winter of 1900.
A female killed near Santa Cruz measured 6 feet 10 inches ;
and a male killed near Lake Argentino, 8 feet 1 inch.
274 MR. H. PRICHARD ON PATAGONIAN MAMMALS, [Apr. 15,
In strong contradistinction to the habit of Felis onca, Felis
concolor puma, when hunting, kills a number of animals from a
flock or herd. To only one of these kills, however, does it return,
and it always makes some pretence of burying the victim singled
out for its meal, throwing up upon the body in many cases
merely a small bunch of thorns. This habit of the Puma is
frequently taken advantage of by the shepherds, who poison the
chosen carcase. The Puma, in ninety cases out of a hundred,
makes its first meal upon the entrails of the victim, or upon the
inside of the thigh by the groin. Another point in connection
with the predatory habits of the Puma is the fact that it will
travel a long distance, even as much as ten or twelve miles, after
killing.
Its method of attack, judging from an examination of its kills,
appears to be to spring upon the shoulders of its quarry and to
break the neck. The destruction wrought by Pumas among
flocks of sheep is immense. One Puma is said to have killed
from a single flock upwards of 100, its total fora night amounting
to 14. Casesare reported of Pumas having attacked horses ; and
sometimes a herd of cows, with their calves, take up the trail of a
Puma witha great deal of lowing, but do not follow it far. Felis
concolor puma usually selects a tempestuous night for its depreda-
tions upon the herds. Authentic instances of their having
attacked man are few. Dr. F. P. Moreno tells me that on the
bank of the River Leona, not far from Lake Viedma, he was
attacked bya Puma. He was walking, wrapped up in a Guanaco
skin capa, and he fancies the animal mistook him for a Guanaco.
The Puma was killed by his companions, and was found to be in
milk. Its cubs, however, were not discovered. Mr. Arenberg,
of the Argentine Boundary Commission, was also attacked by one
of these animals in the neighbourhood of Lake Buenos Aires.
T have no details of the occurrence beyond the fact that he was
wounded in the face. These two instances must be regarded as
exceptional, for the Puma is ordinarily a very cowardly animal,
and many are killed yearly with the bolas or lasso.
The Puma can easily be galloped down, as it rarely runs more
than 300 yardsor a quarter of a mile when pursued on horseback.
It invariably stands at bay with its back to a bush or rock.
Darwin writes that “the Puma is a very silent animal, uttering
no cry even when wounded, and only rarely during the breeding-
season.” In the forests upon the slopes of Mount Buenos Aires
near Lake Argentino, one moonlight night, two Pumas circled
round our camp, and for upwards of an hour kept uttering their
peculiar ery. On no other occasion during our marches, although
Pumas often stampeded the horses and left plain tracks of their
presence close to the camp, did I hear them break silence.
3. Puarson’s Puma. (Felis concolor pearsont.)
On my return from Patagonia, I brought with me a skin of a
Puma, which seemed to me to differ in some essential respects
1902.] MR. H. PRICHARD ON PATAGONIAN MAMMALS. 275
from any known species. Mr. J. G. Miillais, on examining the
skin, agreed with me, and pointed out that it possessed several
characteristics which do not occur in Felis concolor puma. I took
the skin to the Natural History Museum, where Mr. Oldfield
Thomas came to the conclusion that the animal was a sub-species
of Felis concolor puma, and named it Felis concolor pearson.
The chief points of difference between elis concolor puma and
Felis concolor pearsoni are as follows :—The very different general
colour of Felis concolor pearsoni, being reddish fawn instead of
silver-grey; the proportionately very short tail; light instead of
dark colour on the backs of the ears, which are, moreover, more
sharply pointed in the case of Felis concolor pearsoni; and the
absence of dark markings round the digital pads.
Several Gauchos, settlers, and Indians informed me that there
were two kinds of Puma in Patagonia, one being very common,
grey in colour, and very cowardly. The other they described as
rare, much fiercer, of a reddish colour, and somewhat smaller than
the grey common species. Among the 73 Pumas killed by the
English pioneers near Lake Argentino, one, Mr. Cattle told me,
differed very much from the ordinary Puma, and, judging from
the description he gave of it, I have no hesitation in concluding
that it was a specimen of Felis concolor pearsoni.
4, Tue Guanaco. (Lama huanachus.)
(Huanaco of settlers, Argentines, and Chilians; Row of
Tehuelches.)
During the whole course of our travels in Patagonia (save
when in the forests) a day rarely passed without our seeing
Guanacos. They may be met within a few hours’ ride of any
settlement. The range of the Guanaco extends all over the
plains of Patagonia. In my experience they were most numerous
in the Cafadon Davis, in the neighbourhood of Bahia Camerones,
and on the high basaltic tablelands to the south of Lake Buenos
Aires, At the base of the Cordillera, and in some of the river-
valleys under the edge of the mountains, the range of the
Guanaco crosses that of the Huemul. I do not think, however,
that the Guanaco ever enters the forest: although I have seen
them in the open patches amongst the lower wooded parts of the
Cordillera. As the seasons change they move from lower to
higher ground, but these migrations are limited; and a white
Guanaco has been observed year after year in the same neigh-
bourhood. During the time I spent at Lake Argentinc—from
Ist February to 15th May—I saw but few of these animals, for
at that season all the herds migrate to the high pampa. A
herd 300 or 400 strong inhabited the higher plateaus of Mount
Frias.
FitzRoy, in his “ Voyages of the ‘Adventure’ and the ‘ Beagle,’”
writes :—
“Do the Guanacos approach the river to drink when they are
276 MR. H, PRICHARD ON PATAGONIAN MAMMALS. [Apr. 15,
dying? or are the bones and remains of animals eaten by lions or
by Indians? or are they washed together by floods? Certain it
is that they are remarkably numerous near the banks of the
river (Santa Cruz), but not so elsewhere.”
It is true that, although one comes upon skeletons of these
animals upon the pampas, they are not crowded together as they
are in the caiiadones or by the lakes near water. At the edge of
a lagoon at the eastern end of Mystery Plain a great number
of skeletons were to be seen. They extended in a wide track
down the hillside and to the edge of the water. At Lake Viedma
the margins of the lake, near the outflow of the River Leona,
were covered with their skins and bones. The meaning of this I
gathered from Mr. Ernest Cattle. He told me that in the winter
of 1899 enormous numbers of Guanaco sought the Lake Argen-
tino, and died of starvation upon its shores. In the severities of
winter they seek drinking-places where there ave large masses
of water likely to be unfrozen. The few last winters in Pata-
gonia have been so severe as to work great havoc among the
herds of Guanaco.
At nightfall Guanacos gather into close order, a large herd
collecting in a small radius. They seem to choose open spaces in
which to pass the hours of darkness. In moments of danger also
they pack together densely. At the sound of a shot, the outlying
members of a herd will close up and sway their long necks almost
to the ground in unison. I see that Darwin says that Guanaco.
are “generally very wild and wary.” In places where they are
hunted by the Indians this is no doubt the case, but on this point
no law can be laid down. In some districts the Guanaco is very
difficult of approach: in others extremely easy. Their instinct
of curiosity is very largely developed. During our wanderings I
studied the habits of the Guanaco with ever increasing interest.
In cold weather they become extraordinarily tame, and will
permit a man to walk among them as a shepherd walks among
his sheep.
The young are brought forth in the months of October,
November, and the early part of December. In Southern
Patagonia some are born as late as the end of December. During
the period of copulation the bucks fight a good deal. I never
shot an old buck which was not seamed and scarred with the
marks of these contests. When fighting they give vent to loud
squeals of rage; they kick with their fore feet and bite savagely,
mostly at the neck of the antagonist. The marks of these bites
are often deep and long. The ‘skin of the neck is very thick. As
has been noted before, the Guanacos drop all their dung in one
spot, and near these spots their wallows are ordinarily to be found.
I saw an old buck spend a long time over his toilet while his
wives looked on and waited. He would pass nearly half an
hour on his back with his legs in the air, at intervals standing up
to neigh, and then rolling again.
P.Z.$.1902.vol.1 PLXXXI.
H.Grénvold.del. “4y/-ac!
Photogravure by Bale «Danielsson Lt4
OS VEO ROGY Ch TEE aA CO NiO mii Se
\
Poze: LO vole eal
H_Gronvold, del. Photogravure by Bale x Danielsson 1®
OSTEOLOGY OF “FRE Ane ONT OR VES:
be
i
size
P.Z.5.1902,vol.1.P]. XXXUI.
\
map.
H.Gronvold.del. (P 5 Bouin Photogravure by Bale «Danielsson Lt4
Yeas c L
OSTHOUVOEY (OER 2 Ale ONiir On vars
1902.] ON THE OSTEOLOGY OF THE FALCONIFORMES. 277
5. Paracontan Cavy. (Dolichotis patagonica.)
(Called “ Cavy” or “ Hare” indiscriminately by the English
settlers; Liebre by the Chilians and Argentines; Paahi by the
Tehuelches.)
The River Deseado forms the southern limit of the distribution
of the Patagonian Cavy. In 1833, Darwin writes, concerning
the Cavy: ‘“ They are found as far north as the Sierra Tapaleuen
(lat. 37° 30’), and their southern limit is between Port Desire
and San Julian, where there is no change in the nature of the
country.” So far as my experience goes, I never observed a Cavy
after 23rd October, upon which day I counted fourteen upon the
pampa between Lake Musters and the settlement of Colohaupi.
The residents at Colohaupi informed me that that place formed
the southern limit of distribution of the Cavy. It is of course
impossible to lay down an exact line, but I think I am safe in
saying that the range of the Cavy does not extend south of the
46th parallel. This limit is the more remarkable inasmuch as
the country south of lat. 48° does not in any way materially
differ from that over which the Cavy is commonly to be met with.
It is upon patches of dry mud that these animals are most often
to be observed.
6. Lirrnz ARMADILLO. (Dasypus minutus.)
(Pichy of the Chilians and Argentines ; Ano of the Tehuelches.)
This Armadillo is never found south of the River Santa Cruz.
During the four months I spent south of that river I did not see
one, but when, for three days, we crossed to the north bank, we
met with four and killed one. Dasypus minutus is very commor
in the vicinity of the Bahia Camerones. I saw no specimen in
the forests of the Andes, but near Lake Buenos Aires and Lago
Viedma we found them at the foothills.
6. Contributions to the Osteology of Birds.
Part V. Falconiformes’. By W. P. Pycrart, F.Z.8., A.L.S.
[Received March 4, 1902. ]
(Plates XXXI-XXXIIT & text-figures 33-37.)
ConTENTS.
i. Introductory Remarks, p. 277. vii. The Pectoral Limb, p. 306.
ii. The Skull of the Adult, p. 278. viii. The Pelvic Limb, p. 307.
iii. The Vertebral Column, p. 292. ix. Summary, p. 311.
iv. The Ribs, p. 295. x. Key to the Osteology of the
v. The Sternum and Pectoral Girdle, Falconiformes, p. 318.
p- 296. xl. Explanation of the Plates, p. 319.
vi. The Pelvic Girdle, p. 301.
i. IntTRoDUcTORY REMARKS.
The anatomy of the Falconiformes presents many characters in
1 For Part IV. see P. Z. S. 1899, p. 1018.
2 For explanation of the Plates, see p. 319.
278 MR. W. P, PYCRAFT ON THE [Apr. 18,
common with certain of the Grues on the one hand, and of the
Steganopodes and Herodiones on the other.
It will be the aim of the present paper to supplement the
evidence of these affinities which has been so far collected, by a
careful study of the Osteology of the Falconiformes,—a study
which shall embrace comparisons of the skeletons of the forms
not now regarded as allies of this group. By these means it may
be possible to arrive at more definite conclusions concerning the
probable inter-relationship of the undoubtedly allied forms, and of
their phylogeny.
ii, THe SKULL OF THE ADULT,
It seems to be impossible to draw up, in concise terms, any
definition which shall serve us to readily distinguish the skull of
the Falconiformes from any skull of what we are agreed to regard
as the near allies of this group. ‘This is due partly to the fact
that the group contains some very aberrant forms with peculiar
skulls, and partly to the fact that the characters upon which
we are wont to rely for taxonomic purposes present us with many
grades of perfection even amongst the smaller divisions of the
group. Thus, the palate affords a series of gradations between
complete desmognathism and complete schizognathism ; _basi-
pterygoid processes may be very well developed or completely
absent; the lachrymal may be free or fixed; the nares may be
pervious or impervious; and so on.
The best that we can do is to say that no indirectly desmo-
enathous Falconiform skull owes this form of desmognathism to
the intervention of the vomer alone—as in the Cariamide. Again,
no indirectly desmognathous Falconiform skull combines this form
of desmognathism with a large vomer and a large anterior palatal _
vacuity. No Falconiform skull has supra-orbital grooves, or has
the palatines fused in the middle line posteriorly. Finally, all
the skulls of this Order have an ossified orbito-sphenoid.
The resemblances which the skull of the Falconiformes presents,
through certain of its members, to the Steganopodes, Grues, and
Striges will be pointed out and discussed in the following
remarks.
The Occipital Region.
The plane of the occipital foramen varies, from an oblique angle
to one almost parallel with the long axis of the skull. The forms
in which the plane is almost horizontal are those which represent
the higher types—the Eagles, Buzzards, and Falcons. The foramen
in these forms thus looks downwards, instead of downwards and
backwards.
In the Falcons, Buzzards, and Cathartz there is a prominent
cerebellar dome. This is less well-marked in the Vultures and
Eagles, and in Serpentarwus.
The supra-foraminal ridge is fairly well-marked in every
1902.] OSTEOLOGY OF THE FALCONIFORMES. 279
member of the group. It is continued downwards on either side
to form the inner border of the processus ale exoccipitalis inferior
(paroccipital process, azct.).
The lambdoidal ridge is always single and has an undulating
outline. It is continued outwards for some considerable distance,
when it suddenly bifurcates, one branch running directly outwards
and forwards to terminate at the superior angle of the exoccipital
wing, at its junction with the squamosal process; and one directly
downwards and outwards to lose itself in the free border of the
exoccipital wing near its middle. In Serpentarius, Polyboroides,
and the Vultures the upper and outer branch is barely perceptible
or wanting. In Catharte there is a deep notch between the
bifurcation, so that it appears, at first sight, to be wanting. We
may probably regard this as the more primitive form, since
the filling-in of this notch seems to have been to increase and
perfect the tympanic aperture. Beneath the lambdoidal ridge,
and on either side of the cerebellar prominence, lies a well-marked
depression, the supra-occipital fosse. The latter appear to be
best developed in the Falcons. In the Catharte they take the
form of wide channels,
The exoccipital wing is continued downwards into the processus
ale exoccipitalis inferior of Suschkin (paroccipital process, awet.).
These will be found most strongly developed in the Cathartz.
The exoccipital wings are much more developed in the Striges,
where the upper and outer portion of the wing is developed on
either side into a prominent, outstanding flabellum.
The cranial roof (Pl. XXXII.).—The cranial roof is never
marked by supra-orbital grooves.
In Serpentarius (Pl. XXXII. fig. 2) the interorbital region is
widened by long, horizontal, backwardly-directed lachrymal pro-
cesses, the inner borders of which are closely applied to the frontal,
but do not fuse therewith. In the Catharte (Pl. XXXII. fig. 1)
there would appear to have been similar processes, but these have
now fused completely with the frontal, a row of foramina only
indicating their line of junction.
In the Accipitres (Pl. XXXII. fig. 3) these horizontal processes
are free, and project on either side of the head, above the
orbits. In the Buteonide there is a small free plate of bone
attached by igament to the end of the horizontal process—the
superciliary plate. This plate is probably a remnant of the
supra-orbital chain of ossicles such as occurs in some Paleognathe,
e. g. Tinamus, Struthio, and in Psophia among the Neognathe.
Both the horizontal process and the superciliary plate have under-
gone great reduction in the Vultures, Gypohierax alone retaining
a small remnant of the accessory plate. This same reduction of
the horizontal process is also a marked feature in Polyboroides, _
wherein it is almost suppressed; the descending process of the
lachrymal, it may be mentioned in passing, is very long and
slender. The lachrymal of Pandion has undergone still further
degeneration, having fused completely with the frontals and
280 MR. W. P. PYCRAFT ON THE [Apr. 15,
antorbital plate (prefrontal). The horizontal process is practically
suppressed,
The fronto-parietal region in Gypohierax, Aquila, Falco,
Polyborus, and Milvus, the parietal only in Polyboroides, Buteo,
and Halaétus, is marked by a shallow median groove. This
groove is more or less traceable in all the Accipitres save the
Catharte and Serpentarius. It is especially noticeable in the
forms just enumerated. In the Catharte the roof of the skull
presents an evenly rounded surface. The width across the skull
at the fronto-parietal region in no case approaches that of some
Owls, e. g. Bubo, owing to the smaller size of the postorbital
processes.
The fusion of the nasals with the frontals is complete, and
leaves no trace of the line of junction.
The Base of the Skull.
The basitemporal plate in Serpentarius only is visibly thickened
by pneumatic tissue. In the remaining members of this group
it is a thin triangular plate with a slightly concave ventral
surface. Posteriorly it is bounded, in the middle line, by a more
or less well-marked precondylar fossa. It extends outwards on
either side as a wing-like plate to join the inferior wing of the
exoccipital process, in Serpentarius, Kagles, Buzzards, Falcons,
and Vultures, for instance. But in the Osprey this junction with.
the exoccipital—completing the mouth of the tympanic cavity
below—is formed only by a thin bar of bone. In certain Vultures
and in the Catharte the hinder angles of this plate appear to
terminate in a pair of prominent mammillary processes. ‘They
are the dominant features of this region of the plate, and by
their great size have come to lie behind the actual posterior angles.
The two sides of this triangular plate may have sharply defined
free edges, e. g. in Aquila, in which case the Eustachian channels
are open grooves; or they may be partly fused with ossified
connective tissue forming the anterior wall of the recessus
tympanicus anterior, when the grooves are partly closed, e. g.
Serpentarius, Haliaétus, Buteo; or they may fuse throughout
with the inferior border of the wall forming the above-mentioned
recess, leaving only a small Eustachian aperture at the apex of
the triangle, as in the Catharte and Polyboroides, for instance.
In the Catharte the parasphenoidal rostrum immediately above
this aperture is deeply excavated. This is especially marked in
Sarcorhamphus.
The parasphenoidal rostrum may or may not bear basipterygoid
processes. These are largest in Serpentarius, where they lie at
the base of the rostrum. In the Catharte they may be either
short and broad, as in Catharistes and Gypagus, or very slender
and seated further forward on the rostrum, as in Pseudogryphus
and Sarcorhamphus (Condor). ‘This greater slenderness evidently
marks the first stages in their decay. In a skull of Sarcorhamphus
1902. ] OSTEOLOGY OF THE FALCONIFORMES, | 281
in the collection of the British Museum the left process is smaller
than the right, and both almost fail of their purpose, not only
in this species but throughout the genus, They present within
the Order every degree of degeneration, from the prominent pillars
of Serpentarius to the most minute prickles.
Traces of the anterior basicranial fontanelle are often present.
The Lateral Surface of the Cranium, (Plate XX XIII. figs, 1-8.)
The tympanic cavity attains its greatest size in the Falcons.
It may be studied in its more primitive form in such examples
as Polyboroides, Pandion, Pernis, and the Vultures, The lateral
occipital wings which bound this cavity behind in Polyboroides
are but feebly developed. The free border, on either side,
terminates in a blunt and low prominence behind and slightly
above the level of the processus zygomaticus, and marks at the
same time the lower and hindmost extent of the temporal fossa,
In Pandion the free border of this wing, which is still feebly
developed, slopes obliquely backwards and upwards to the level
of the processus articularis squamosum, then turns sharply
forwards to terminate in the process itself. Pernis resembles
Pandion in this respect, but neither the backward slope nor the
forward angle is so marked, In Serpentarius and the Vultures,
e.g. Neophron, Gypohierax, and G'ypaétus, the free edge of this
wing is of greater extent, and projects as a slightly raised ridge
beyond the zygomatic process. [In Gyps the form of the lateral
occipital wing is drawn backwards and downwards to terminate
in a prominent processus ale exoccipitalis inferior (paroccipital
process), and thus in this respect differs from Meophron, where
the wing is more feebly developed and slopes from before back-
wards and upwards.| In Buteo the ridge laterad of the zygomatic
process is much more prominent. In Haliaétus and Aquila it has
increased still more. In the Falcons it forms a thin laminate
plate, rising upwards to the level of the base of the zygomatic
process of the squamosal, the free edge of which is turned
forwards. In the development of this portion of the exoccipital
it bears a strong resemblance to the Striges, e. g. Bubo.
The roof of this cavity is formed by the under surface of the
processus zygomaticus squamosi. Its floor, in part by the
lateral occipital wing, and in part by the ossification of tissue
extending between this wing and the external angles of the basi-
temporal plate. In front it is bounded by the quadrate.
Within this cavity, in the dried skull, are two large apertures
which may be considered separately. The first hes near the
roof of the cavity and separates the squamosal and otic articular
surfaces of the quadrate. The aperture leads upwards and
backwards, so as to form a space between the anterior vertical
and horizontal canals of the internal ear and the lateral occipital
wing. This is the recessus tympanicus superior—the temporal
recess of my earlier papers.
The second and lower aperture is divided from the first by the
Proc. Zoot, Soc.—1902, Vou, I, No, XIX, ug
282 MR. W. P. PYCRAFT ON THE [ Apr. 15,
articular surface for the otic head of the quadrate. Within it
lie several foramina and the mouths of two pneumatic cavities.
The foramina are the foramen ovale and the foramen rotundum,
the foramen for the 7th nerve, and the foramina of the sinus
petrosus. The pneumatic apertures are, as already stated, two
in number. The first, and largest, opens into the mouth of the
tympanic cavity at its antero-ventral angle. It may be traced
inwards, and forwards, as a tubular recess leading into the para-
sphenoidal rostrum, terminating in the pituitary region. The
anterior wall of this recess—the recessus tympani anterior—is
formed in its larger exterior portion, as is shown by Suschkin *,
by ossification of connective tissue extending in the young
skull between the alisphenoid above and the basisphenoid below.
When the Eustachian grooves are closed, they form two additional
and much smaller apertures, opening one on either side, into the
tympanic cavity, below the mouth of this anterior tympanic recess.
The second of these apertures is that of the posterior tympanic
recess. It is very small and not easily seen. If carefully looked
for, it will be found as a small hole, lying caudad of the foramen
ovale and foramen rotunda. It leads into a small pneumatic
cavity lying below the horizontal semicircular canal, and between
this and the inferior border of the lateral cccipital wing.
The separation of the squamosal and otic articular surfaces
for the quadrate by the aperture of the superior tympanic
recess is a point of some interest. In the Paleognathe the
recessus tympani superior isrepresented by a shallow cavity, lying
behind the articulation for the head of the quadrate. The roof
of the cavity is pierced by numerous small pneumatic apertures
leading from a mass of diploé lying between, and above, the
horizontal semicircular canal and the brain-case, The external
wall of this diploid tissue is formed by the parietal and lateral
occipital bones. In some Dinornithide there is a small aperture
connected with this mass of pneumatic tissue lying in front of
the articulation for the quadrate. In the Neognathe the aperture
of the superior tympanic recess lies—in Steganopodes, Tubinares,
and Sphenisci, for instance—in front of the quadrate articular
surface. But in the Accipitres and Striges, for instance, it
would seem that the separate anterior and posterior apertures,
lying on either side of the articular surface for the quadrate, i
the Dinornithide, have here become confluent, and now form one
large aperture dividing the squamosal and otic articular surfaces
for the quadrate far from one another. In the Sphenisci, by
reason of the great depth of the temporal fossa lying immediately
above, the greater part of this recess has become suppressed, only
the lower end now remaining. In the Pygopodes the relatively
greater depth of the temporal fossa, coupled with a general and
marked tendency for the suppression of pneumatic tissue through-
out the skeleton, have combined to obliterate the superior
tympanic recess altogether.
1 “Yur Morphologie des Vogelskelets.” Nouveaux Mém. de la Soe. Imp. des
Naturalistes, 1899,
1902. | OSTEOLOGY OF THE FALCONIFORMES. 283
The Squamosal Prominence——In the more primitive forms,
such as Serpentarius, Polyboroides, and Pernis, the squamosal
prominence is very feebly developed. As usual, its inferior
surface affords a glenoid cavity for the articulation of the squamosal
head of the quadrate. Mesiad of this glenoid cavity is the aperture
of the superior tympanic recess. The zygomatic process is very
small; in Serpentarius it is wanting.
The squamosal prominence is seen at its best in the Falconide
(including Polyborus), where it projects conspicuously from the
skull-wall. The zygomatic process is excessively developed in
Polyborus, where it hangs downwards, eaves-fashion, over the
quadrate. In addition to this, is a second and equally large
process depending from the antero-lateral angle of the lateral
occipital wing, at its junction with the zygomatic process.
The postero-inferior angle of the squamosal, in Polyboroides
and in the Falcons, develops a large, downwardly-directed and
pointed spike—the processus articularis squamosi. Between this
and the zygomatic process just described, the squamosal head of
the quadrate is tightly grasped. In Serpentarius, the Eagles,
Pernis, Pandion, Buteo, Circus, Gypactos, and many of the true
Vultures, for instance, this articular process is but feebly
developed. In Cathartze it is wanting.
The temporal fosse are for the most part but shallow, linguiform
depressions in the parietal region of the skull, and which never
extend backwards to meet the mid-dorsal line. In Serpentarius
they are exceedingly shallow and scarcely rise above the level of
the base of the postorbital process. They are best developed in the
Falcons and Polyborus, where they extend inward to within a
a short distance of the cerebellar prominence.
The trigeminal foramen is completely isolated, lying far removed
from the mouth of the superior tympanic recess, and considerably
in front of a line drawn transversely through the skull across the
squamosal head of the quadrate.
The orbits are large. In Serpentarius they are protected from
above by wide overhanging ledges formed by the frontal bones,
behind, and by large, horizontal, backwardly-directed, flattened
plates developed by the lachrymal, infront. These, one on either
side, fit closely to the frontal. In the higher Accipitres the width
of the interorbital region of the frontals is much less, the free
edge forming a deep hollow. In this case the horizontal process
of the lachrymal is left as an isolated spur projecting on either
side of the roof of the skull and overhanging the orbits. In the
majority of such cases, the length of the spur is increased by
the addition of a separate scale-shaped ossicle—the superciliary
ossicle. This is attached to the lachrymal by connective tissue,
In the Falcons and Vultures it fuses with the lachrymal. In the
Catharte the supra-orbital limb of the lachrymal is suppressed,
the lachrymal itself fusing completely with the frontal.
The orbito-sphenoid is completely ossified in all the Falconi-
formes,
19*
284 MR. W. P. PYCRAFT ON THE [Apr. 15,
The interorbital septum is pierced only in the lower members of
the various groups of Falconiformes. For instance, in Serpentarius
and Cathartes, in Pandion and Pernis, Polyborides, Gypohierax,
and the lower Vultures and Eagles.
The Kthmoidal Region.—The mesethmoid, as in Neognathee
generally, is, as it were, obliquely truncated at its anterior end,
so that its free edge slopes upwards and forwards, This is due
probably to the shortening of the parasphenoidal rostrum. This
shortening process 1s very conspicuous in the Accipitres. It
rarely, if ever, extends forwards beyond the level of the lachrymals
in any Neognathe. In the Paleognathe this rostrum extends
forwards for a considerable distance in front of a line drawn
transversely through the skull across the lachrymals. The
mesethmoid extends backwards, as in all other birds, to fuse with
the orbital plate of the frontal, and the orbito- and presphenoid,
and the parasphenoidal rostrum below; thus forming the inter-
orbital septum referred to above. From the neighbourhood of
its truncated anterior border springs, on either side, a more or
less extensive wing-like process—the prefrontal or antorbital
late.
b The prefrontal is somewhat feebly developed in Serpentarius,
Polyboroides, and Pernis, for instance. In Catharte and in the
Falcons it is much larger; in the latter it extends outwards to
afford a more or less extensive lateral support for the lachrymal.
In the former, the lachrymal and prefrontal relations become still
more intimate, since they fuse one with another.
The preorbital region of the mesethmoid expands dorsad into
a broad horizontal plate underlying the anterior ends of the
frontals, the nasals and the nasal processes of the premaxillary.
In the Vultures, Eagles, and Catharte markedly, and in the
majority of the Accipitres to a less extent, the free edges of
the horizontal aliethmoidal plate turn downwards and inwards,
and finally backwards, to join the prefrontal; thus forming an
ossified olfactory chamber. This is most perfectly developed in
the Cathartz, where the chamber is of very considerable extent,
recalling that of the Tubinares. In Serpentarius, the Falconide,
and Buteonide, this ossified olfactory chamber is extremely
reduced. In the two latter forms perhaps the great development
of the prefrontal may be regarded as filling the place of the
ossifications of the horizontal plate.
The olfactory nerve, in leaving the skull, generally travels along
a groove in the dorsal border of that portion of the mesethmoid
which forms the interorbital septum; sometimes this groove is
covered in by the ossification of connective tissue, e. g. Cathartze.
I would revert once again to the comparison between the
olfactory chamber of Cathartee and that of the Tubinares. In
the former the free edge of the horizontal aliethmoidal plate
turns downwards on either side mesiad of. the lachrymal, so as to
leave a considerable space between itself and the lachrymal, to
form the Harderian fossa, Furthermore, it would seem that the
1902.] OSTEOLOGY OF 'HE FALCONIFORMES. 985
aliethmoidal extensions of this plate extend backwards so as
to form a complete olfactory chamber leading backwards directly,
tunnel-wise, into the brain-cavity, through the apertures for the
olfactory nerve.
In the Tubinares we have a very similar olfactory chamber,
but of far greater size. Thisisdue to the fact that the prefrontal,
which is enormous, arises so far back that the interorbital septum
is reduced to a small ring of bone surrounding the interorbital
fenestra. Furthermore, the backward position of the prefrontal
has involved it in the formation of the lower lateral segment of
the aperture for the olfactory nerve, which may also be legitimately
described asenormous. The upper lateral segment of this circular
aperture is formed by the downturned edge of the horizontal
aliethmoidal plate. Normally, in Neognathe, the olfactory nerve
leaves by a small foramen pierced through the free edge of the
orbital plate of the frontal and the perpendicular plate of the mes-
ethmoid, runs in a groove along the dorsal border of the plate,
and thence gains the olfactory chamber. In the instance we
have just examined, the olfactory aperture opens directly into the
chamber.
It is possible that the conditions which obtain in the Tubinares
may represent a primitive stage, the olfactory chamber having
been. pushed forwards by the great development of the orbits,
demanded by the increasing size of the eye. In Apteryx, where
the eyes are very small, the olfactory cavity is in close juxta-
position with the brain-case. The fact that the chamber is of still
greater relative size, and very complicated internally, only indicates
afurther specialization of the primitive condition. In the Palzo-
gnathe we may trace several stages in the forward shifting of
the olfactory chamber correlated with increased size of the eyes
and orbits; as a study of the skulls of Dinornis, Struthio,
Dromeus, Rhea, and Tinamus will show.
The development of the olfactory cavity is a point which will
evidently repay further investigation.
The nasal septum is divided by a considerable gap, or cleft,
from the mesethmoid. This cleft, the cranio-facial fissure, is widest
ventrally ; the mesethmoid and nasal septum being in contact
immediately below the nasal processes of the premaxilla. In the
Catharte alone among the Falconiformes, the external nares are
pervious, only the proximal portion of the nasal septum being
present. This, in Pseudogryphus californianus, extends forwards
for a short distance to encroach upon the external narial aperture,
and is pierced by a round fenestra. This posterior portion of the
nasal septum, in Catharte, expands to form a broad base which,
extending outwards on either side, joins the widely separated
maxillo-palatine processes, and so converts a schizo- into an in-
directly desmognathous palate. In many of the true Vultures,
e. g. Gyps, by the ossification of the alinasal ectoethmoidal wall,
the nostril, in the dried skull, is of the same shape as in life.
The olfactory region of the nasal labyrinth, in Gyps, is provided
286 MR. W. P. PYCRAFT ON THE [ Apr. 15,
with an ossification of the basis of the choncha media which
projects into the cavity from the outer wall. In the true Falcons
the ossification of the aliethmoid causes the anterior nares to be
round in form as in life, in Polyboroides crescentic. In both, the
aperture displays within its mouth a small median bony papilla—
the papilla of the conche vestibulum of Suschkin, the alinasal
turbinal of W. K. Parker. The details of the structure of the
cartilaginous nasal labyrinth and its ossifications have been
exhaustively worked out and beautifully illustrated by Dr. Suschkin
in his masterly monograph on the skull of Zinnaneulus.
The Cranial Cavity.—The metencephalic fossa of the Falconiformes
is more basin-shaped in the smaller than in the larger forms.
Moreover, in these smaller forms the anterior region of the fossa
is tilted upwards and backwards so as to form an acute angle with
the long axis of the skull. The trigeminal foramen, in Falco for
instance, is sharply cut off from the mesencephalic fossa by an
overhanging ledge of bone. The orbito-nasal foramen also les
immediately under this ledge. In the Vultures, the upper ledge
of bone overshadowing these apertures is less extensive, so that
they come to lie partly in the floor of the mesencephalic fossa.
The apertures of the vagus and internal auditory meatus do not
offer any very important points of difference for comment.
The cerebellar fossa is sharply defined and variable in relative
proportions, being, for instance, larger in Yalco than in Vuliwr or
Circus. The floccular fossa is, in Falco, cordiform, and apparently
rather shallower than in other forms.
The mesencephalic fossa is very sharply defined. It is banded
above by a broad overhanging ledge formed by the tentorial ridge.
The distinctness of its inferior border is blurred, in Vuléwr, by
the apertures of the trigeminal and orbito-nasal nerves, which lie
in the floor of the fossa. In Falco these are shut off from this
fossa by a bony shelf, and open inward into the dorso-lateral
border of the metencephalic fossa.
The piturctary fossa is deep and tubular, and passes almost
vertically downwards. The hinder boundary of this fossa, the
dorsum selle, forms a narrow ridge passing forwards and upwards
to terminate above the oculo-motor foramen. The anterior
border of the fossa is bounded by the transverse pre-pituitary ridge,
which passes forwards into a narrow, sometimes triangular, optic
platform, on either side of which lie the optic foramina. The
optic platform is continued upwards, forwards, and backwards
into the pre-optic ridge which may be traced, in Circus for instance,
on either side into the tentorial ridge. In the majority of other
forms, probably, it disappears before reaching this, e. g. Vuléwr,
Falco.
The oculo-motor lies immediately caudad and ventrad of the
optic foramen. It is continued backwards in a‘rather wide groove
into the dorsum selle. Below and anterior to this foramen is
the aperture for the internal ophthalmic artery—when this is
separate. It opens into the pituitary fossa some distance from the
1902.| OSTEOLOGY OF THE FALCONIFORMES, 287
aperture for the internal carotid. The pathetic lies dorsad of the
oculo-motor foramen, and may be continued backwards and down-
wards by a groove terminating above the orbito-nasal foramen.
The cerebral fosse do not lie altogether in front of, but rather
exhibit a tendency to overlap the cerebellar fosse. The lateral
extension of the cerebral fossee is very marked. The fosse are
conspicuously depressed dorso-ventrally. This is most noticeably
so in Falco: the distance between the point where the right and
left tentorial ridges converge to join the median bony falx and
the prominent triangular bony boss which bounds the mesen-
cephalic fossa posteriorly being considerably less than in Vultur.
The olfactory fosse are represented by a small: median pit
leading forwards into two narrow apertures for the optic nerves.
The Premacxilla.
The premawilla, in all the Falconiformes, is more or less dis-
tinctly hooked at the tip. In length and stoutness it varies.
In the Falcons and Polyborus it is short and wide and much
hooked. The tomium, in Falcons, is notched, or rather is
provided with a distinct “tooth.” The palatal surface in both
Falcons and Polyborus is extensive and marked by a well-defined
median ridge, passing backwards in the maxillo-palatine processes.
This ridge is faintly represented in Serpentarius.
In many Vultures, the palatal surface of the premaxilla is more
or less deeply hollowed (Plate XXXI. fig. 5); the excavation
being most noticeable in the larger forms. In others, where
the breadth is slender, the palatal surface is but slight, passing
rapidly backwards into a groove filled by the ventral border of the
nasal septum.
In the majority of the Falconiformes, the palatal surface of
the premaxilla resembles that of the smaller Vultures—is slight,
bifurcate, and filled by the ventral surface of the nasal septum.
The Cathartze differ markedly from all the other Falconiformes
in the form of the palatal surface of the premaxilla. In these
last the body of the premaxilla is very highly pneumatic,
a section thereof revealing a mass of cancellated tissue between
two dense plates of bone. This increase of pneumatic tissue
brings the palatal surface down close, or very near to, the
level of the tomium. In the Cathartze this pneumatic tissue is
almost wanting, so that the palatal surface comes to form a
vaulted chamber. The resemblance in the structure of the pre-
maxilla to the Tubinares is very striking.
The fusion of the nasal processes of the premaxilla with one
another and with the nasals is very complete in all the Falconi-
formes.
In Microhierax the beak articulates with the skull by means of
a fronto-nasal hinge as in Parrots and some other birds.
-In the Catharte the fused lachrymal and frontal combine to
288 MR. W. P. PYCRAFT ON THE [Apr. 15,
form, with the nasal bones, a more or less perfect fronto-nasal
hinge, see also p. 291. There is no transverse hinge as in
Microhierax or in Parrots. .
The Masillo-Jugal Arch.
The maxilla is indistinguishably fused with the premaxilla.
The maxillo-palatine processes have slowly increased in size so as
to convert what -was originally a schizognathous into a desmo-
gnathous palate. Many intermediate stages in the course of this
transformation have been preserved and will be discussed forth-
with.
In the study of the transition of the palate from the schizo-
gnathous to the desmognathous type, we will commence with
Hlanus: this form being regarded by Dr. Suschkin as exhibiting
more primitive characters than any other Accipitrine bird.
In Hanus then (Pl. XX XT. fig. 2) the maxillo-palatine pro-
cesses turn inwards in the form of a pair of small, very pneumatic,
hamulate processes. Although they hem-in the vomer on either
side, they do not extend so far inwards as to touch it. Distad
of the vomer they are hidden by the palatines, and thus leave
a large median palatal vacuity exposing the nasal septum above
and lying in the middle line. The maxillo-palatine processes
themselves are pierced, laterad of the palatines, and between
these and the tomium, bya large foramen. ‘The palate, however,
of Hlanus is not, I think, to be regarded as primitive, but
specialized (see p. 313). Cireaétus (Pl. XX XI. fig. 4) is also schizo-
gnathous, and probably represents the intermediate type of palate
which has given rise, by specialization, in one direction to the
palate such as is found in Hlanus and the Catharte, and in the
other to such as obtains in the Vultures and Falconide.
In Aquila (e. g. A. chrysaétos, A. audasz), Spilornis, and Thras-
aétus harpyia we find the maxillo-palatines greatly increased in
size antero-posteriorly, and extending inwards mesiad of the
palatines: so much so as to almost completely fill up the space
between them. The nasal septum is now visible only through a
narrow palatine clink. This increase in bulk has now caused the
maxillo-palatines so nearly to approach one another, that they
almost, if not quite, touch. But they are yet distinct, and the
palate must accordingly be regarded as schizognathous.
In Serpentarius they touch throughout the greater part of their
length, concealing the nasal septum (Pl. XXX. fig. 7).
Indirect desmognathism, brought about by the downward
extension of the nasal septum, obtains in the majority of Accipitres,
and can best be studied in such forms as Hlanoides, Pandion,
Vultures. In Gypaétus and Neophron, for example, the maxillo-
palatines do not meet in the middle line, and but for the nasal
septum would be bounded anteriorly by a large palatal vacuity
as in Cathartes.
Direct desmognathism is perhaps only found in Falconide.
1902. ] OSTEOLOGY OF THE FALCONIFORMES. 289
In all the desmognathous palates the maxillo-palatine processes
remain free posteriorly.
In the Catharte we have a palate of quite peculiar type
(Pl. XXXTI. fig. 1). The maxillo-palatines, as in Hlanus, are
relatively small. But whilst, in Hlanus, it will be remembered,
they were of a spongy texture, in Catharte they are represented
by a shell-like lamina with its concavity downwards. The external
lateral border of this lamina is anchylosed with the nasal. The
antrum is represented by a small cavity at the extreme anterior
extremity of the plate. These shell-like lamine fail to meet in the
middle line; nevertheless the palate must be regarded as of the
indirect desmognathous type on account of the fact that the nasal
septum expands ventrally into a horizontal plate, fusing on either
side with a pair of strap-shaped lamine arising from the inner
dorsal border of the maxillo-palatine processes (=the anterior
septo-maxillary spur, Parker). Distad of the maxillo-palatines and
nasal septum is a iarge anterior palatal vacuity roofed only by the
nasal processes of the premaxilla. This region of the jaw recalls
that of the Tubinares. But the palate of the Tubinares differs
in the absence of an ossified nasal septum and the great size of
the vomer.
The jugal is wanting from the quadrato-jugal arch in the
Falcons. Itis certainly present in Accipiter, and perhaps in other
Accipitres.
In some examples of MWierofalco the quadrato-jugal bar is
connected with the maxilla by a distinct joint. The maxilla con-
tributes towards this articulation a backwardly directed, columnar
bony process, which immediately overhangs the extreme posterior
angle of the tomium. Seen from below, the articulation is
transverse in form. From above, it is concealed by a spike of
bone from the quadrato-jugal, which projects forwards into the
mouth of the antrum. The sporadic formation of a joint in this
position seems to me a point of some considerable interest.
The Vomer, Palatine, and Pterygoid.
The vomer in the Accipitres is never a very large bone. It is
blade-shaped when completely developed, and sometimes is slightly
bifurcated at its extreme posterior end—seen from the ventral
surface. Its base is received between the anterior borders of the
dorsal lamin of the posterior expanded end of the palatine:
sometimes fusing therewith, e. g. Serpentarius. In the Falcons
the vomer terminates in a bilobate expansion which rests upon
the fused maxillo-palatine processes, between their free posterior
projections. In Polyboruws and Milvago this expansion is barely
perceptible. In Milwus (e.g. WM. ater), Haliastur, Pernis, and
Buteo (e. g. B. jakal) the vomer is long and slender, and anchy-
loses with the maxillo-palatines. In Aquilinw the vomer extends
far forwards between the maxillo-palatines, but remains perfectly
distinct therefrom. ;
290 MR, W. P. PYCRAFT ON THE [| Apr. 15,
In the Vultures the vomer appears to be wanting, save in
Gypactus, where it is large. In Neophron and Gypohierax vestiges
remain, and it may be that these traces are lost in maceration in
other skeletons. In the remaining Accipitres it is either vestigial
or wanting.
The palatine perhaps more nearly resembles that of Carrama
than any other. It extends forwards as far as or slightly beyond
the level of the external narial apertures, running beneath the
maxillo-palatine process. It increases gradually in width from
before backwards, reaching its maximum expansion at a point
immediately below the prefrontals. From this point backwards
it comes into relation with the parasphenoidal rostrum; the palatine
of the right and left sides combining to form a narrow, very
shallow trench, gliding up and down the rostrum with the move-
ments of the facial apparatus. That portion which is connected
with the rostrum is somewhat scroll-like in form, the free anterior
edge of the scroll passing downwards and forwards into the mesial
border of that portion of the palatine which is continued forwards
to terminate below the maxillo-palatines as a rod- or rather strap-
shaped blade. Seen from below, it would seem as if the scroll-
shaped region of the expanded palatine plate were caused by a
grooving out of substance of the palatine itself, causing a deep
median trough, bounded on either side by a sharply-defined ridge
representing the mesial border of the palatine, e.g. Hlanus and
Serpentarius. In other Accipitres, in varying degrees, this ridge
projects downwards and forms a more or less conspicuous keel.
This keel is especially strongly marked in the Falconide.
The pterygoid in the lower forms, e. g. Serpentarius, Cathartes,
bears an articular surface for the basipterygoid process. In the
Accipitres it forms either a cylindrical or twisted rod-shaped
bone. The adult palato-pterygoid articulation is of the typical
Neognathine type in all but Pandion. In Pandion the anterior
end of the pterygoid is Y-shaped, the external fork of the Y¥
articulating with the external postero-lateral angle of the palatine.
This is, so far as I am aware, a quite unique feature; but never-
theless a quite secondary character, and in no way comparable to
the forked pterygoid of Apteryx. .
The maxillo-jugal bar in the Cathartz furnishes some interesting
matter for study, in certain peculiar features connected with its
anterior end. These features may be traced through a series of
stages gradually increasing in complexity, and associated with a
gradually developing fronto-nasal hinge.
In Cathartes (C. awra) this bar, when it reaches the middle of
the lachrymo-nasal fossa, splits into a long inferior dorso-ventrally
depressed lamella and a superior shorter rod-shaped portion,
which extends forwards to within a short distance of the maxillary
process of the premaxilla. [In one skull (1186 6) in the Museum
Collection, on the left side, the terminal portion of this maxillary
process, it should be mentioned, is segmented off to form a very
small separate triangular ossicle. |
1902.} OSTEOLOGY OF THE FALCONIFORMES. 291
In Catharistes the fission of the bar takes place much further
forward than in Cathartes, and the short superior segment is
abruptly truncated and roughly articulates with the aforesaid
maxillary process of the premaxilla.
In Gypagus this inferior limb remains much as in Cathartes
and Catharistes, but the superior limb now forms a shorter but
laterally compressed lamina with a hollow anterior border fitting
into the rounded posterior border of the maxillary process of the
premaxilla.
In Pseudogryphus and Sarcorhamphus the short, laterally com-
pressed upper limb now forms a spatulate process, working in
opposition to a shallow depression in the maxillary process of the
premaxilla.
The gradual development of these peculiar articulations seems
to have gone hand in hand with the development of a nasal
hinge, formed by a deep notch hollowed out of the anterior
border of the lachrymal, for the reception of a “ tooth-shaped ”
process borne by the outer border of the nasal. This forms the
hinge; the necessary movement of the beak upon the skull is
afforded by the elasticity of the nasal and premaxillary processes
lying mediad of the right and left notches. This hinge is most
perfect in Gypagus, Pseudogryphus, and Sarcorhamphus. In
Catharistes the lachrymal is only barely perceptibly notched, A
further study into the origin of structures like the present would
prove profitable. Are they to be ascribed to kinetogenesis ?
The composition of the quadrato-jugal bar, as revealed by an
immature skull of Cathartes, is interesting, and chiefly on account
of the large share which is borne by the maxilla. This, together
with the jugal, extends backwards as a long slender bar to
within a short distance of the quadrate articulation. The
jugal terminates anteriorly to meet, and sometimes articulate
with, the maxillary process of the premaxilla, and apparently in
Pseudogryphus and Sarcorhamphus, for instance, becomes much
broadened to make the articulation the more complete. The
quadrato-jugal appears to terminate at about the middle of the
inner aspect of the bar. <A slightly older skull of Serpentarius,
but still showing traces of sutures, agrees very closely with what
has just been described in Cathartes. The great backward exten-
sion of the maxilla appears to be a Neognathine character.
The Me andible.
The mandible of the Accipitres is truncated posteriorly, and
has a very long internal angular process. It very closely
resembles that of the Bubonine section of the Striges, so much
so as to require very careful discrimination. In comparing
mandibles of these two groups, it will be found that among the
Accipitres the two rami form a less open angle one with another,
and the ventral aspect of the internal angular process is not
marked by a median ridge. The lateral vacuity in the mandible
299 MR. W. P. PYCRAFT ON THE [ Apr. 15,
of the Accipitres is wanting save in the Falconide, wherein
it 1s conspicuous. It differs, however, from that of the Striges
(1) in that the coronoid extends forwards along its whole
length, and (2) in that there is a small foramen lying immedi-
ately behind and above the vacuity for the passage of the
mandibular branch of the trigeminal. Furthermore, the external
lateral border of the articulare develops a strong upward and
backwardly directed hook-like process forming a deep notch
between itself and the extreme postero-lateral angle of the jaw.
In the Striges, the Bubonide have the coronoid terminating in
a spike-like fashion near the middle of the lateral mandibular
vacuity. In the Strigide the vacuity is much smaller; and the
coronoid terminates in front of it, in a truncated and spatulate
process. The V-shaped angle of the jaw is similar to that of the
Accipitres; but the internal angular process is relatively more
feeble, and the inferior border of the jaw is sinuous, whilst in the
Accipitres it is nearly straight.
The Hyoid.
The basihyal is partly ossified; short ceratohyals are also
represented. The basibranchial has a diamond-shaped body
continued backwards into a long style, representing the 2nd
basibranchial (urohyal). The ceratobranchials are long. The
epibranchials are more than half as long as the ceratobranchials.
il. THE VERTEBRAL CoLuUMN.
All the presyrisacral vertebre are heterocelous. In all, save
the Falconide and Polyboride, the thoracic vertebre are free.
In the two families just mentioned all the thoracic vertebra are
ankylosed save the penultimate, which is free. Herpetotheres is
the only exception to this rule, having all the thoracics free.
The cervical vertebre are relatively thick and short, and have
in the larger forms a somewhat swollen and massive appearance.
The ankylosed cervical ribs, which are present in all but the
atlas and axis, never extend backwards as far as the end of
the centrum. The anterior lateral border of each is produced
upwards into a flattened plate, forming the outer wall of the
vertebrarterial canal. This pleurapophysial Jamella is short
antero-posteriorly, its posterior border never extending as far
backwards as the middle of the centrum. The dorsal border of
this lamella never unites with the centrum to enclose a fenestra
opening immediately behind the posterior zygapophyses, as in
Cariama for instance. Neural spines take the form of low median
tubercles: in the smaller forms these are but feebly developed.
In the 2nd to 4th vertebree the neural spines are moderately long
and columnar, especially in the Falcons; the posterior angles of
the 2nd and 3rd turn upwards into blunt tubercular hyper-
apophyses. In Pandion the neural spines are obsolete, but the
1902. | OSTEOLOGY OF THE FALCONIFORMES. 293
hyperapophyses are particularly well developed. The 2nd to 4th
vertebree bear hypapophyses: beyond these they are replaced by
catapophyses, which near the cervico-thoracic vertebra again
give place to hypapophyses. These catapophyses never coalesce
to form a carotid canal. In many of the larger Accipitres the
cervical vertebree from say the 5th to the 8th have the neural
plate deeply excised posteriorly, so that the postzygapophyses
appear, each as an articular facet at the extremity of a long beam.
The ventral surface of the centrum is either flattened or slightly
grooved.
There is a very close similarity between the cervical vertebre
of the Accipitres and the Striges: so close is this resemblance
that the one is hardly distinguishable from the other. The chief
differences appear to be in the fact that the cervical ribs of the
Striges are relatively slightly longer and the catapophyses some-
what more sharply defined. The pleurosteal lamella is also
somewhat more band-like and sharply defined. The hyper-
apophyses of the axis are abruptly truncated in the Owl, tubercular
in the Accipitres. For the rest, the differences are not greater
than those which normally obtain between species or genera, at
most,
The cervici-thoracic vertebrae may be two or three in number.
The thoracic vertebre, as already remarked, are free save in
the Falconidz and Polyborids. When free, they may be dis-
tinguished from those of the Strigide in that the neural spines
are relatively lower, wider antero-posteriorly, and not markedly
inclined forward. Hypapophyses in the Strigide do not extend
beyond the second vertebra: in the Accipitres with free vertebree
to the third. In the Strigide there may bea large pneumatic
aperture between the articular surfaces for the capitulum and
tuberculum, and a second caudad of this. As in the Accipitres
so in the Striges, each transverse process sends forward and
backward a long slender spike from its extreme lateral border ;
each spike overlaps similar spikes from the vertebra next in front
and behind it. The neural spines may also send backwards from
the upper border a pair of short spike-like processes, to embrace
the neural spine immediately behind it.
In Serpentarius the thoracic vertebree, from the 2nd to the 5th,
are pierced by a large pneumatic foramen, opening at the base of
the neural spine immediately behind the anterior zygapophysis.
This leads into an extensive chamber, excavated out of the
vertebral tissue and extending down to the spinal cord, being
separated therefrom only by a thin plate of bone. Other
pneumatic apertures pierce the lateral walls of the neural tube,
and the centrum below this.
In the majority of the Accipitres, the pneumatic apertures of
the thoracic vertebre are restricted to a single opening at the
base, and caudad of the transverse process and immediately in
front of the base of the postzygapophysis.
Jn the Catharte the apertures are three in number, and lie on
294 MR. W. P. PYCRAFT ON THE [ Apr. 15,
either side of, rather than between, the capitular and tubercular
glenoid sur faces,
In the Falconide, excepting only Herpetotheres, and in Poly-
boridee, as already remarked, the thoracie vertebre 1—4 are anky-
losed into a single mass. But this mass also includes the last
cervico-thoracic vertebra, so that the whole is made up of five
vertebra. The penultimate thoracic vertebra is free, the ultimate
is fused with the synsacrum ; so that this free vertebra serves as a
hinge in the middle of the back. In some examples of Polyborws
the antepenultimate vertebra may show traces of its former
existence. Such specimens are probably not quite adult.
It is probable that the last two cervico-thoracic vertebra have
not long been transferred to the cervical series, for the dorsal
segment of the ribs belonging thereto has as yet undergone no
shortening. Those of that vertebra which has ankylosed with
the thoracic still retain their uncinates.
In the Faleonide and Polyboride only one, and in other
Accipitres two thoracic vertebre appear to enter into the syn-
sacrum,
The synsacrum includes from 12 to 14 vertebrae. In Serpen-
tarius it is composed as follows :—2 thoracic, 4 lumbar or cruro-
sacral, 2 lumbo-sacral or ischiadic, 2 sacral, and 4 caudal vertebree.
The anterior renal fossa (fossa ischiadica) in Serpentarius is
very lofty and narrow ; when seen from the ventral aspect, recalling
that of Cariama. The posterior renal fossa is long and narrow.
This last fossa is bounded by a very broad planwm anale, and this
appears to be peculiar to Serpentarius, amongst the Falconiformes.
In the Accipitres, the form and size of these fossee vary much,
as is indicated by the following illustrations. It is impossible,
however, to do more than indicate the general nature of the
form of these fossee, for they vary in details in almost every
species. The anterior renal fossa forms a narrow and lofty
chamber, longer than the posterior fossa, in Busarellus, Circaétus,
Haliaétus, Haliastur, Lophaétus, Gypohierax, and Antenor.
It is narrow, lofty, and shorter than the posterior fossa in
Parabuteo, Aquila, and Spizaétus. It is wide, lofty, and longer than
the posterior fossa in Malco, Chimachima ; wider, lofty, and shorter
in Hlanoides and Polyborus. In Pandion both these fosse are
exceptionally large, and of the type seen in Hlanoides. In
Spizaétus the posterior renal fossa has reached the minimum limit
of reduction, including but a single caudal vertebra.
It will thus be seen that the form of these fosse is of little or
no value for taxohomic purposes other than the identification of
species. The lofty and narrow type appears to belong to the
more specialized, the wide and shallow to the more primitive
members of the respective groups to which they belong.
In Pandion, there are three and sometimes four lumbar or
cruro-sacral vertebre, and these are more sharply defined than in
the other Accipitres. The lumbar possess well-developed dorso-
lateral processes. The sacrals are, in the skeleton, indistinguish-
1902.] OSTEOLOGY OF THE FALCONIFORMES. 295
able from the postsacral (caudal) vertebrae by reason of the long
slender ventro-lateral processes of these last, which in no respect
differ in appearance from the sacral ribs. Thus the posterior
renal fossa (fossa pudendalis), which is large, is cut up into a
number of separate compartments.
There is a certain resemblance between the synsacra of Pandion
and Cathartes. But the two may readily be distinguished. In
Pandion the lateral iliac fossa (fossa lwmbaris) is peculiar on
account of the great size of the ventri-lateral processes, the outer
ends of which are fused one to another to form a bony bar, upon
which the preacetabular ilium rests. The dorsi-lateral processes
forming the roof of the anterior and posterior renal fosse give
rise to a large, flat, imperforate plate with convex free border S,
showing a distinet suture-line betwixt them and the postace-
tabular ilia. The intervertebral foramina are almost completely
obliterated.
In Cathartes the ventri-lateral processes of the lateral iliac fossa
are less conspicuously developed. The roof of the anterior and
posterior renal fossee is perforated by large intervertebral foramina ;
and the lateral edges of the plate forming the roof of this fossa
is deeply hollowed both before and behind the sacral vertebre.
There are 6 to 8 postsynsacral vertebre (free caudals) including
the pygostyle. The anterior of these vertebre bear vestiges of
the anterior zygapophyses, which gradually decrease in size from
before backwards.
In Serpentarius the neural spines are swollen and _ bifid, the
transverse processes are pierced at the base by comparatively
large pneumatic apertures. The last 3 or 4 vertebre bear bifid
hypapophyses. The free caudals of Gypaétus resemble those of
Serpentarius but are non-pneumatic, In the larger Eagles the
transverse processes are much expanded and very wide. The
number of caudal vertebre included in the synsacrum appears to
vary between | (Spizaétus) and 4 (Serpentarius). In the Falconide
the pygostyle bears a pair of accessory plate-like vesicles, attached
by their hinder border by ligament to its inferior angle, and pro-
jecting forwards as a horizontal plate, Intercentra occur.
iv. THE Riss.
The cervical ribs extend from the third vertebra backwards,
and are ankylosed with their respective vertebre. They are
relatively longest in Serpentarius. 'The free, so-called cervico;
thoracic belong more correctly to the thoracic series.
The thoracic ribs range from 5 (Pseudogyps, Polyboroides) to 8
pais. The 8th pair, however, is never complete: often, as in
Spizaétus for instance, only the sternal segment is present, fixed
by membrane to the 7th pair of sternal ribs. Sometimes the
thoracic and sternal segments are widely separated by loss of the
intermediate ossified tissue, as in a case of Aquila chrysaétos.
Whilst in other cases the thoracic and sternal segments are
296 MR. W. P. PYCRAFT ON THE [Apr. 15, ,
normally related, yet the latter fails to reach the sternum. In
some cases the last pair of sternal ribs may be represented only
by vestiges fused with the last fractional pair.
The uncinates vary considerably in their development. In the
Polyborine and the Falcons the uncinates are long and slender ;
but in the remaining Accipitres the base of attachment to the rib
has increased enormously, extending often nearly as far down as
the articulation with the sternal segment. In the Catharte, and
in some species of Aquila, the inferior angle of this base is
produced into a short spine. Generally the uncinates do not
extend backwards beyond the level of the rib next behind. This
is exceeded in Hlanoides, and in Pandion (in P. carolinensis) they
extend each to the third rib. In Serpentarius the uncinates are
very degenerate, losing themselves in the rib, forming thereon
nothing but a broad irregular plate. The uncinate of the first
thoracic rib, however, is less degenerate, projecting. distinctly
backwards.
The preacetabular ilium overlaps from two to three pairs of
ribs.
v. THE STERNUM AND PECTORAL GIRDLE.
Perhaps the most striking features of the sternum of Falconi-
formes are the great size of the corpus sterni and the relatively
slight development of the carina. Amongst the different
members of the group, however, much variation occurs, in the
relative proportions in the length and breadth of the sternal
plate, the development of the keel, the position of the coracoid
grooves and articular surfaces for the sternal ribs, and the de-
velopment of notches or fenestra on the metasternum.
In Serpentarius (text-fig. 33, p. 297) the sternum is distinctly
Ciconiine in character, not only on account of the great size and
shape of the carina, but also in the fact that the latter affords a
broad articular surface for the furcula. There is one peculiarity
about the keel of Serpentariws, however, and this lies in the fact
that the carina, swollen by pneumatic tissue, passes gradually into
the corpus sterni, This last, it should be mentioned, is pointed
posteriorly, instead of notched as in the Storks.
The sternum of the Catharte is peculiar in that the greatest
curvature of the free margin of the carina is near its middle, and
in that the keel extends backwards to the extreme posterior end
of the sternal plate. The corpus sterni is notched posteriorly,
and in Cathartes the posterior lateral processes are fenestrated.
The coracoid grooves are shallow and broad, curving abruptly
upwards and inwards to the middle line, not overlapping. The
spina externa and interna are both wanting. The linea aspera
for the origin of the subclavius extends backwards to within a
short distance of the posterior end of the sternal plate: thus
_ extending further back than in any other Falconiformes.
In the Falcons only is there a distinct spina interna; in Herpe-
totheres it is very broad and deeply hollowed; the spina externa
1902.] OSTEOLOGY OF THE FALCONIFORMES. 297
is present in all the Accipitres. In the Falcons only does the
anterior angle of the carina project forwards as far as the level
of the spina externa. Again, whilst in nearly all the other
Accipitres the sternal ribs attach themselves along the whole
outer border of the anterior lateral processes, in the Falcons
the distal third is ribless. In the larger Falcons the sternal
plate is fenestrated posteriorly, but in the small forms—WVelierax,
Poliohierax, Microhierax—it is notched. In Falcons, the coracoid
Text-fig. 33.
Car
Left lateral aspect of the Sternum and Shoulder-girdle of Serpentarius serpentavius,
showing the articulation of the furcula with the carina.
ac. = acrocoracoid. car. = carina.
cor. = coraco:d. J. = furcula.
cs. = corpus sterni. sr. = sternal rib.
H, = hypocleideum.
grooves overlap one another, and the spina externa is relatively
long and pointed. The articular surfaces for the sternal ribs
extend backwards as far as the middle of the sternal plate.
The sternum of the Polyborinw resembles that of the Falcons,
but the spina interna is shelf-like instead of spiny, and the
anterior border of the keel does not project so far forward.
In the Buteonide (Suschkin) the spina interna is not developed.
The spina externa varies much in size. Generally, it nay be
described as a short stout tubercle; but in Accipiter 1t becomes a
Proo. Zoou. Soc.—1902, Vou. I. No. XX. 20
298 MR. W. P. PYCRAFT ON THE [Apr. 15,
long spine, as in Falco, The sternum varies much both m its
relative length and the form of its posterior border. Thus, this
last may be either notched or fenestrated, or both, or it may be
entire. As these are points which may vary in members of the
same genus, they are of comparative little value save as specific
characters.
The sternum with the relatively largest carina of this group is
that of Pandion. The carina generally has its anterior border
sloped backwards, but in Acczpiter this border turns forwards and
upwards, thus greatly increasing the length of the keel. The
carina varies much in its development, generally passing insensibly
into a sternal plate before reaching its hinder border, but in
Accipiter and Pernis, for instance, the carina is carried back
almost to the extreme free border of the sternal plate.
The sternum of the Accipitres resembles that of the Striges.
In many cases this resemblance is rather close. The following
characters will be found useful in determining between sterna
belonging to these two very different groups. The posterior
border of the sternum in the Striges is never entire and never
fenestrated, but always notched. With the exception of the
sterna belonging to the Strigide, and the sternum of Huhua
orientalis of the Bubonide, there are two pairs of notches. The
single pair of notches of H. orientalis are of great size. They lie
on either side of the metasternum, and extending forwards to
beyond the level of the middle of the posterior lateral process,
cause the sternum of this bird to closely resemble that of
Microhierax. The great difference in size, however, renders any
possibility of confusion on account of this resemblance impossible.
The single pair of notches of the Strigide resemble those of some
Accipitres, e.g. Hlanoides, in that they are very shallow, so that
the posterior lateral processes pass almost insensibly into the
metasternum, being divided therefrom only by asinuousline. But
the processes are relatively much longer in the Strigide than in
the Accipitres, and the sternum is narrower. Furthermore, the
sternum of the Owls of this group may be distinguished by the fact
that the spina externa is obsolete. The anierior lateral processes
in all the Owls are small. The articular surfaces for the sternal
ribs never extend beyond the middle of the sternal plate, rarely so
far as this. There is no spina interna.
The coracoid presents two forms :—(1) That which obtains in
the Falconid only, wherein there is a distinct hook-like pro-
coracoid process curving downwards till it touches the furcula ;
and (2) that in which this process is wanting.
In the Falconide we can distinguish the coracoid of the
Faleones from that of the Polybori; “inasmuch as in the former
there is no trace of the foramen supracoracoideum in the dried
skeleton, this being bounded internally by membrane only. In
the Polybori this foramen is present.
. Whether the foramen in the coracoid of the remaining Accipitr es
is really the foramen supracoracoideum, or a foramen pierced
1902.] OSTEOLOGY OF THE FALCONIFORMES, 299
through the coracoid proper, for the passage of this nerve, is a
point which can only be determined by an appeal to embryo and
nestling specimens. Possibly it will be found that the foramen
in the Polybori is also a coracoid foramen.
The coracoid is relatively of medium length in the Hanlegunuky
but proportionately longer in the little Microhier ax.
In the Buteonide it is relatively short and stout, and very
broad at the base, the distance across the latter region, at its
widest part, being equal to the length of the shaft from the base
to the articulation with the scapula.
In Elanoides the processus lateralis basalis is very conspicuous,
more so than in any other member of the Order,
Text-fig. 34,
Left lateral aspect of the Sternum and Shoulder-girdle of Aquila rapax. The furcula
does not articulate with the carina; and the latter terminates before reaching
the end of the sternal plate.
Letters as in text-fig. 33.
The acrocoracoid process 1s large in both sections of the Order,
and bears a facet for the articulation of the furcula—except only
in Serpentarius, wherein this facet is wanting. The absence of
a procoracoid process is sufficient to distinguish the coracoid of
Serpentarius from that of the Storks.
The coracoid of the Cathartze, like that of the Falcons, is
relatively long.
The coracoid of the Accipitres closely resembles that of the
Striges. In the latter group, however, the procoracoid process is
always present; on this account, dageetore. the coracoid of the
Buteonine section of the Accipitres is always distinguishable.
For the same reason—the lack of the procoracoid process—the
coracoids of the Serpentari and Catharte are also easily dis-
90%
300 MR. W. P. PYCRAFT ON THE * [ Apr. 15,
tinguishable. With the Falconid, however, the case is different ;
and it becomes a matter of nice discrimination to tell the coracoid
of the Falcones, or Polybori, from that of one of the Striges. The
Falcones may be distinguished by the absence of a supracoracoid
foramen; but in the Polybori and Striges this foramen is present,
and almost identical in size and position. The coracoid of the
former can, indeed, scarcely be distinguished from that of the
latter. In the Owls apparently the base of the coracoid is com-
paratively deeply grooved to fit the dorsal coracoid lip of the
sternum ; whilst in the Polybori, what corresponds to the dorsal
lip of the groove in the Owls is represented only by a low and
incomplete ridge; lastly, in the Owls the linea aspera for the
coraco-brachialis posterior ‘generally lies near the extreme lateral
border of the coracoid shaft; in the Polybori it lies nearer the
middle line.
The scapula does not afford much matter for comment of any
systematic value. The acromial process in the Falconine is
pneumatic and very broad, so much so as to project considerably
into the foramen triossewm, thereby considerably diminishing its
size. The acromion may be produced conspicuously forwards,
and this is particularly well seen in Meophron, where it con-
stitutes the greater part of the inner wall of the foramen triossewm.
The furcula in the Accipitres is remarkable for its great size
and strength. It is U-shaped, with the limbs set wide apart,
this latter feature being especially noticeable in the Buteonide.
In Serpentarius only does the furcula articulate with the
carina (text-fig. 33, p. 297), as in the Storks, Furthermore, as
we have already pointed out, Serpentarius agrees with the Storks
in that the furcula does not develop a facet for articulation with
the coracoid. It bears a hypocleideum of considerable size, directed
downwards, and presenting a keel-like border for articulation with
the carina.
In the Catharte only are the distal extremities pierced by
a pneumatic foramen. This is very large and opens in the outer
aspect of each limb, immediately behind the acrocoracoid.
In the Falconide the hypocleideum is vestigial or wanting.
In Microhierax the furcula is not perceptibly bent upon itself. as
in the larger members, nor are the limbs so wide apart. The
length of the limbs of the fureula in Mierohierax is relatively
greater than in any other members of the order, since they equal
the length of the carina sternt.
In the Buteonide the limbs of the furcula, as we have already
remarked, are set widely apart, and are of great breadth at their
distal ends; especially is this the case in such forms as G'ypaétus,
Haliaétus, and Aquila. The hypocleideum is vestigial or wanting.
It appears to be largest in Aguila and Pandion. In some
cases, Elanus for example, the proximal ends of the furcula are
very slender, and in this particular the furcula resembles that of
the Striges. The furcula of the Accipitres is more unlike that
of the Striges than is the case with the sternum or coracoid, as
1902.] OSTEOLOGY OF THE FALCONIFORMES. 301
we have already shown. With the exception of Serpentarius,
the furcula never articulates with the carina in Falconiformes.
In the Striges, on the contrary, the furcula is always attached
to the carina. Furthermore it is a much more slender bone
than in the Falconiformes, and not bent upon itself.
vi. THe Petyvic GIRDLE.
The pelvic girdle of the Falconiformes, through the more
aberrant members of the group, bears resemblances on the one
hand to that of the Gruide, and on the other to that of the
Ciconiide ; and, through the more specialized forms, to the Striges.
The innominates are never free.
Serpentarius presents several Gruine characters, the most
noticeable of which are the pocket-like cavities (iliac recesses) of
the postacetabular ilium, and the general contour of the dorsal
aspect of the pelvis as a whole.
The pelvis of Serpentarius is, however, distinguishable from
the similar Gruine and Ciconiine pelves by the great height of
the supra-trochanteric process, and the enormous size of the ilio-
ischiadic foramen. Furthermore, the ischium terminates poste-
riorly in a rounded or rather conical border projecting beyond the
postacetabular ilium ; whilst the pubis, which is long and slender,
sends up a conical process immediately below the projecting
extremity of the ischium, which serves more or less effectually to
close the obturator fissure posteriorly.
The preacetabular ilia meet one another in the mid-dorsal
' line, and there is no trace of the suture between the postace-
tabular ilium and the transverse processes of the synsacrum.
The synsacral fosse lying between the neural spines and the
postacetabular ilium are roofed over by a thin plate of bone.
The obturator fissure is not separated from the foramen.
The pectineal process is wanting, not only in Serpentarius but
in all the pelves herein described.
In the Catharte the pelvic girdle is, externally, distinctly
Ciconiine in character. The resemblances are especially notice-
able in the pelves of Psewdogryphus (text-fig. 35, p. 302), Catharisies
(text-fig. 36, p. 303), and Gypagus.
The presence of iliac pockets, however, at once distinguishes
these pelves from those of the Ciconie. Another Stork-like
feature is the deep notch in the hinder border of the innominate,
marking the division between the now-fused ilium and ischium.
In Sarcorhamphus, Pseudogryphus (text-fig. 35, p. 302), and
Gypagus the inferior limb of this notch is produced backwards
for a very considerable distance beyond the postacetabular illum
to form a long spine. In the degree of development, and in the
position of the supra-trochanteric process, the innominate of the
Catharte is Gruine.
The pelvis of Cathartes differs markedly from that of the other
genera in this: That whilst in the genera Just discussed the
302 MR. W. P. PYCRAFT ON THE [ Apr. 15,
preacetabular ilia rise above the synsacrum to meet one another
in the middle line; in Cathartes they are widely separated one
Text-fig. 35.
AUS oe
Dorsal aspect of the Pelvis of Pseudogryphus californianus, showing the Ciconiine
character of the pelvic girdle. The preacetabular ilia rise above the ridge
formed by the neural arches of the vertebra, and the external intertransverse
sacral foramina are almost completely filled up.
acet.=acetabulum. p.=pubis.
ant.tr.=antitrochanter. p.-a.il.=preacetabular ilium.
él.-is.2.=1lio-ischiadic notch. pt.-a.il.=postacetabular ilium.
from another, and do not rise above the synsacrum. The
peculiar form of this pelvis bearsa striking resemblance to that of
1902. ] OSTEOLOGY OF THE FALCONIFORMES. 303
certain Steganopodes. The resemblance is perplexing, and would
seem to suggest that it is from this more primitive stock that the
Falconiformes, in common with the Ciconie, have been derived :
the Ciconiine characters of the Falconiform skeleton, already
alluded to, being homoplastic. On the whole, however, I feel, at
present, inclined to adopt the Gruine origin of the Falconiformes,
suggested by Beddard. Cathartes and Catharistes agree in that
the backwardly produced spine of the ischium does not project far
beyond the postacetabular ilium, The iliac pockets of the Catharte
are conspicuous by their absence.
Text-fig. 36.
Wh — wee & Cr
Yip
pale |
> i) Any
0) NET
iG a
\ \
bir pf A glee
i Yj 7) hy Ye i j
\ Wg - Woe aN
Dorsal aspect of the Pelvis of Cathartes aura.
This is probably a more primitive type of pelvis than that of Pseudogryphus, and
recalls that of Fregata or Phaethon. Note that the preacetabular ilia are
widely separated, lying on either side of the broad ridge formed by the vertebral
neural spines, and that the intertransverse sacral foramina are large and
numerous.
Additional letters.
int.for.=intertransyerse sacral foramina.
2.S.=neural spines.
We now turn to the pelvic girdle of the Accipitres, Here
304 MR. W. P. PYCRAFT ON THE [ Apr. 15,
we meet with a type of pelvis found elsewhere only amongst
the Striges, Its chief characteristic is the great deflection and
shortening which the postacetabular ilium has undergone, and
this is more marked in the Accipitrine innominate. The
innominate bone of the Owls can, however, it would seem, be
readily distinguished by the fact that the ischium is invariably
produced backwards into a more or less slender spine resting
on the pubis.
In its general conformation the pelvic girdle of the Falconi-
formes, after the elimination of the Gypogeranide and Cathartz, is
very uniform: modifications from the type are very considerable.
Some of the more noticeable, however, may profitably be com-
mented upon here. The most conspicuous departure from the
type is found,in the pelvis of Pandion (text-fig. 37, p. 305),
which is remarkably broad. This great breadth is due partly to the
length of the outstanding transverse processes of the synsacrum,
which support the roof of the anterior and posterior renal fossee ;
and partly to the exceptionally broad dorsal plane of the post-
acetabular ilium., The preacetabular ilium is sharply truncated
anteriorly, and is widely separated from its fellow of the opposite
side: so much so that a tubular aperture—the canalis dleo-
lumbalis—is left at the pomt where the mesial curved border
leaves the synsacrum, The glenoid surface of the antitrochanter
is oblong in shape and curved backwards. The ilio-ischiadic
foramen is of great size. The obturator foramen is also very
large, and closed posteriorly by the approximation of the pubis
to the inferior border of the ischium, which, we may remark,
turns abruptly upwards at its hinder end. The pubis is long,
and develops a ecrescentic plate the cephalad segment of which
fits closely to the upturned ischial border; beyond this it is con-
tinued backwards for a considerable distance as a rod-shaped bar
nearly meeting its fellow of the opposite side,
The pelvis of Pandion (text-fig. 37, p. 305) is an exaggeration
of the typical Accipitvine pelvic girdle, which may be very well re-
presented by such a form as is presented in Polyborus for example.
Herein the pre- and postacetabular ilia are about equal in length.
The former is a moderately broad concavo-convex plate directed
outwards and downwards, meeting its fellow of the opposite side
in the middle line, and having its superior border accentuated by
a sharply-defined outstanding crest, which, in some forms, as in
Haliaétus and Parabuteo for example, becomes still more strongly
uccentuated, forming an almost shelf-like projection. This border
is continued backwards to terminate in an overhanging supra-
trochanteric process. The postacetabular ilium expands into
broad dorsal plates, terminating somewhat abruptly some distance
from the end of the ischium, with which itis fused. In Accipiter
and Hlanuws, and still more markedly in Polyboroides, the dorsal
border of the ischium turns sharply inwards so that the dorsal
plate of the ilium forms a relatively enormous ledge overhanging
a deep cavern passing forwards into the obturator foramen, In
1902. ] OSTEOLOGY OF THE FALCONIFORMES. 305
Aquila, Thrasaétus, Haliaétus, Circaétus, for example, the post-ilia
pass Insensibly backward into the ischium, instead of abruptly.
Text-fig. 37.
Dorsal aspect of the Pelvis of Pandion haliaétus.
As in the pelvis of Cathartes, the preacetabular ilia fail to rise above the neural
spines of the vertebre. A pair cf canales ileo-lumbales are present, but most
of the intertransverse sacral foramina have become filled up. The great breadth
of the pelvis, as shown in text-fig. 36, p. 303, is caused by the unusual length of
the sacral ribs and transverse processes.
In the majority of Accipitres the pre- is longer, sometimes
nearly twice as long, than the postacetabular ilium, which is
markedly deflected.
The ischiwm is generally truncated posteriorly, but in some,
as in Thrasaétus and Aquila for example, the posterior end is
hastate.
The pubis is generally long and slender, and at the level
of the end of the ischium turns abruptly inwards towards the
middle line nearly meeting its fellow of the opposite side. In
Thrasaéttus, Lophaétus, Parabuteo, it is vestigial, only the proximal
end. remaining. This terminates immediately behind the ob-
turator foramen, and serves to close it, often by fusion with the.
306 MR. W. P. PYCRAFT ON THE [ Apr. 15,
ischium. In many forms, as in some species of Faleo, HZenpe-
totheres, Elanoides, Elanus, Spilornis, Accipiter, the median
portion of the pubis has disappeared, the proximal portion fusing
with the ischium, immediately behind the obturator foramen,
whilst the distal (hinder) ends are attached to the posterior
border of the ischium. Thus several grades in the degeneration
of this bone are presented. Sometimes the two portions are
connected by a very slender thread of bone.
The iliac recess in Serpentarius is spacious, and extends back-
wards into a pocket-shaped cavity, as in Rails. This pocket is
wanting, or but feebly developed, in the other Falconiformes,
except Pandion, but it is interesting to note that it is universally
present in the Striges.
vil. THe Pecroran Limp.
Not only is there little variation in the form of the wing
between the different members which are included in the present
group; but there is also a strong resemblance between the wing
of the Faleoniformes and that of the Grues on the one hand, and
the Storks on the other: the resemblance to the Grues being
especially marked.
The following characters displayed by the emerus will help in
distinguishing between the wings of these three groups :—In the
Falconiformes and Grues the scar marking the insertion of the
pectoral muscle runs along the free border of the triangular pectoral
crest from the tuberculum externus till it reaches the summit of the
triangle, where it expands into a broad elliptical space occupying
the lower limb, and, in the Catharte, extending on to the shaft.
In the Ciconie the distal extremity of this sear takes the form of
a strongly-raised, lmguiform plate which lies partly on the shaft of
the humerus and partly at the base of the distal limb of the crest ;
instead of occupying nearly the whole crest. The humerus of the
Falconiformes may be distinguished from that of the Grues in
that, in the former, the subtrochanterie fossa is larger, the distal
extremity of the humerus is much wider, and its ulnar tuberosity
much more prominent.
The coraco-humeral groove appears to be markedly developed
only in Pandion. In other forms it is either indicated only by a
very shallow groove, or by two depressions. The head of the
humerus in the Falconiformes is more compressed from palmar to
dorsal surface than in the Grues. The supra-condylar depression
for the brachialis inferior varies in its development. In Pandion,
for example, it is deep with gently sloping sides; in Gypagus the
floor is flat and oval and is bordered by a well-defined rim imperfect
distally: in Falco and <Accipiter again it is barely traceable;
in Serpentarius it is an oval depression placed rather high up in
the shaft. A small tubercle only, represents the ectepicondylar
process. The form of the pectoral crest varies. In Serpentarius,
Pandion, and Falcons-it is sharply triangular, and in Pandion
1902. | OSSLOLOGY OF THE FALCONIVORMES, 307
the proximal border of this triangle is deeply hollowed. In
Accipiter the distal border terminates abruptly. The pectoral
crest is very long, extends far down the shaft, and is semicircular
rather than triangular,
The relative proportions of the arm, forearm, and manus vary.
Generally the manus is the shortest segment, and the forearm the
longest. In Valeoninw and Llonus, for exarmple, the manus is
the longest and the humerus the shortest. But the forearm is
always longer than the humerus.
The forcurm, as a rule, shows only faint traces of tubercles
marking the attachment of the quill-feathers, In Catharte,
however, these tubercles are very strongly developed, thus re-
calling the ulna of the Ciconi#, The radiva of the Falconiformes
is more nearly cylindrical than in the Grues.
The manus, as we have already remarked, may be longer than
the humerus, it never exceeds the forearm, The third metacarpal
is slender, laterally compressed, and, save in Serpenturiuas, is
attached only by its extreme proximal and distal ends. The
manus may on this account be more or less readily distinguished
from that of the Ciconisz or the Grues. In the members of both
these groups the proximal end fuses with the second metacarpal
distad of a line drawn transversely through the metacarpus at
the level of the articulation for the pollex. The distal end of
metacarpal III, in the Falconiformes, Serpentarius excepted,
resembles that of the Ciconix, and differs, like the latter, from
that of the Gruide and Serpentarius, wherein the inter-
metacarpal space is lessened by the deposition of bony tissue.
It is extremely difficult to distinguish the manus of the smaller
Accipitves from that of the Striges of similar size, Perhaps the
only character which will hold good for a large series of com-
parisons is that afforded by the point of fusion of the proximal
end of metacarpal III. In the Owls this takes place distad of a
line drawn through the metacarpus at the articulation for the
pollex, as in the Ciconiz and Grues,
The phalanges do not call for any special comment. The pollex
bears a small daw. The postaxial border of the only phalanx of
digit III. has the middle of the postaxial border raised into a
small tubercle,
viii. Tue Petvic Lime.
The pelvic limb in a]] the Falconiformes, save Serpentariua, is
shorter than the pectoral, But whilst, as in Polyboroides, the two
limbs are nearly equal in length, in Serpentoriue the difference is
very considerable.
The femur, in all, is pneumatic, a pneumatic foramen opening
on the dorsal aspect of the femur, mesiad of the base of the great
trochanter—as in the Ciconiz. The popliteal fossa is but feebly
developed. The shaft is moderately long, and cylindrical,
The tibio-torsus is in all more or less dorso-yentrally depressed,
308 MR. W. P. PYCRAFI ON THE [ Apr. 15,
and is slightly bowed forwards. The cnemial crests are not
largely developed.
In Serpentarius they form. prominent outstanding blades, not
extending down the shaft but strictly confined to its proximal —
extremity. The ectocnemial crest is directed outwards, standing
at right angles to the entocnemial. In so far as the direction of
this crest is concerned, it agrees with the remaining Falconi-
formes, but in its greater size it stands alone. Furthermore, the
shaft of the tibio-tarsus is peculiar, in that it is perfectly straight
and almost cylindrical, not bowed forwards and depressed as in the
other Falconiformes. The tibio-tarsus thus much more closely
resembles that of the Storks. Additional Stork-like characters
are found at the distal end of the shaft in the great breadth of
the extensor bridge, and in that the lateral borders of the
posterior trochlear surface are produced backwards and upwards
into a pair of prominent ridges. But there is no intercondylar
tubercle, and no depression below the extensor bridge. The
trochlez are not laterally compressed, but separated by a wide and
deep gorge. ‘There is a conspicuous entocondylar tubercle. The
fibular crest is but feebly developed, but the fibula extends down
to the lower third of the shaft.
The tibio-tarsus of the Catharte is much more Accipitrine in
character. The fibular crest is prominent. The extensor bridge
is wide, oblique, placed nearer the inner border of the shaft, and
somewhat more superficial than in the other Falconiformes.
In the Falconide and Buteonidz there is not much difference,
save in minor points, some of which, however, are extremely
helpful for diagnostic purposes.
Take the Falconide for example. In Herpetotheres, immediately
above the outer tibial condyle, and laterad of the extensor bridge,
is a shallow pit. This, in Polyborws, becomes pierced by a
small foramen opening inwards into the extensor groove. In
the remaining Falcons (Herpetotheres being the exception) and
in Milwago this small foramen has increased to an aperture as
large as that below the extensor bridge. Thus there appears to
be two extensor bridges, one above each condyle. Apart from
size, the tibio-tarsus of Herpetotheres can easily be distinguished
from that of Polyborus by the fact that in the latter the bridge is
wider and the intercondylar gorge is narrower and deeper,
In the Buteonide the most aberrant tibio-tarsus is that of
Pandion. One of its most striking characters is the extreme
depth and width of the extensor groove, which is so deep that it
is saved from perforating the shaft only by a very delicate plate
of bone. The fibula extends to within a short distance of the
tarsal segment, and is very large. The fibular ridge is placed
somewhat lower down the shaft than usual. The distal end of
the shaft, above the extensor bridge, is wider than across the
condyles, which are laterally compressed.
It is a point of considerable interest to notice that the extensor
groove in the Striges is barely perceptible in the Barn-Owls,
1902.] OSTEOLOGY OF THE FALCONIFORMES, 309
deep in the remaining forms, whilst the ossified extensor bridge
is conspicuous by its absence. The absence of this bridge should
surely be as valuable a piece of evidence that Pandion is not
related to the Striges, as is the presence of an extensor bridge
on the tarso-metatarsus to show that it is so related! Further-
more, we may remark that this tarso-metatarsal bridge is wanting
in the Barn-Owls, so that the value of this character is still
further weakened.
The tibio-tarsus of Polyboroides is, in its way, almost as re-
markable as that of Pandion. It is long and almost of the same
thickness throughout. As in Pandion, the width of the shaft
immediately above the distal condyles is greater than across the
condyles themselves. This is probably due to the fusion with
the shaft of the distal end of the fibula which extends down to
the tarsal elements—though, as just indicated, fused with the
shaft. The extensor bridge is placed transversely across the shaft.
The tibial condyles are but feebly developed, the ectocondyle
barely projecting beyond the shaft. The intercondylar gorge is
wide.
The lower third of the shaft of the larger Accipitres is dorso-
ventrally depressed. There is often an additional and well-defined
fibular bridge. The second lies some distance below the first, and
affords attachment for the distal end of the fibula. The cnemial
crests are relatively but slightly developed. In <Agquwila, perhaps,
they reach the maximum development, the entocnemial crest being
unusually strong. In Thrasaétus the ecto- and entocnemial con-
dyles are conspicuously wide apart, causing the inner border of
the shaft to have a very marked curve.
The fibula, as a rule, tapers to a fine point, and terminates near
the lower third of the tibial shaft, sometimes fusing therewith.
In Pandion, Polyboroides, and Pernis only is it of almost uni-
form thickness and continued downwards as far as the tarsal
elements.
The tarso-metatarsus varies extremely, both in its relative
length and in the development of bony matter for the mechanical
requirements of the limb,
In Serpentarius only is the tarso-metatarsus as long as the
tibial shaft. In Accipiter, however, it is very nearly so. Generally
it is shorter than the femur.
The hypotarsus is simple in all save Pandion and Pernis,
wherein it forms a tube.
The distal trochlexw, save in Serpentarius and Catharte and in, 3c pace
Leptodon, are all on the same level. In the two first-mentioned
exceptions the middle trochlea is produced somewhat beyond the
level of the others. In Leptodon the inner trochlea is the longest,
the middle slightly shorter, and the outer shorter still, so that an
obliquely sloping series is formed. The plane of the trochlex
forms a slight and regular curve, except in Pandion, in which this
curve is very strong; and Polyboroides, in which the 2nd and 3rd
trochlee lie close together, whilst the Ist is bent downwards so
310 MR. W. P. PYCRAFT ON THE | Apr. 15,
that the dorsum of the trochlea lies almost behind the level of
the 2nd and 3rd. The regularity of the curve is thus broken.
Usually the outer and inner trochlee are produced backwards
into spur-like processes, that of the outer trochlea being directed
backwards, whilst the inner slopes obliquely outwards and back-
wards away from the shaft. In some few cases, as in Leptodon
for example, these spurs are barely perceptible.
In Sepentarius, the Catharte, and the Falconide the hypotarsus
takes the form of a strong median keel. But whereas in the two
former the keel is grooved equally on either side, and terminates
above in a more or less quadrangular table; in the latter it is
much more deeply grooved externally, and terminates above as a
thin vertical plate with a flanged free border. The front of the
tarso-metatarsus in Serpentarius and Cathartz is deeply grooved.
In the former are two distinct raised surfaces for the tibialis
anticus, in the latter only one. Immediately above the surface
for attachment of the tibialis anticus are two foramina, large in
the Catharte, small in Serpentarius.
In the Falconide the anterior tarso-metatarsal groove is shallow,
the foramina are small, and there is but a single raised process
for the tibialis anticus, which lies on the inner side of the shaft.
In the other Accipitres the hypotarsus takes the form of two
more or less prominent spurs separated by a wide groove. The
inner spur is generally somewhere near the middle line of the shaft,
but in Hlanus it springs from the inner border of the shaft.
The shaft of the tarso-metatarsus is very variable in form. In
Serpentarius it is long and cylindrical, but with the inner aspect
grooved, but in the majority of the Accipitres it is more or
less dorso-ventrally flattened and twisted into long and broad
ledges and plates; thus adding exceedingly to its power.
These features are most marked in Thrasaétus, which in section
is seen to be almost =9-shaped. The same is true of Aquila,
Haliaétus, Parabuteo, Busarellus, Antenor, Urubitinga, and
FHelotarsus, for example, but to a lesser extent. In forms such
as Gypohierax and G'ypaétus the shaft is much less markedly
modified, andthis accords well with their habits. Polyboroides
has a peculiar shaft. This, as we have previously remarked, is
long, much flattened antero-posteriorly, and deeply grooved be-
hind: its outer border is flattened out into a relatively broad
plate, so that in section the shaft resembles that of Thrasaétus
without the lateral torsion.
The phalanges are also characteristic. In Serpentarius the
ungual phalanges are large but not remarkably so. Ph. 1, 2 of
digit IT. are subequal; ph. 2, 3, digit III. are shorter than ph. 1;
ph. 3 and 4, dig. IV. are much reduced.
In Cathartz ph. 1 of the hallux is very long; the phalanges of
digits II., III. are also relatively much longer than in Serpen-
tarius or the Accipitres. The phalanges 2, 3 of dig. IV. are also
longer, relatively, than in other Falconiformes.
In the Faleonide ph. 1 of dig. IT. is considerably shorter than
1902. | OSTEOLOGY OF THE FALCONIFORMES. 311
ph. 2, but not so much as in the other Buteonine. Similarly
ph. 2 of dig. III. and ph. 2, 3 of dig. IV. are relatively much
longer than in the latter group.
In the Buteonide ph. 1 of dig. IT. is very short and often fused
with ph. 2, as in Halaétus for example: ph. 2, dig. III. is
generally much shorter than ph. 1 and 3: ph. 3, 4, dig. IV. are
always very short. In Hlanus ph. 3 is reduced to a vestigial
condition.
In Accipiter ph. 2, dig. ITI. is not shortened: in Hlanus it is
exceedingly so. Again, in Accipiter ph. 2, 3, dig. IV., though
obviously shortened, are not nearly so much so as is usually the
case among the Accipitres. The numerous variations of this
character are useful generic characters, and will be found in the
keys which it is proposed to add to this paper.
It is interesting to note that the foot of the Striges in the
matter of proportionate lengths of the phalanges resembles the
Buteonine section of the Accipitres.
ix. SUMMARY.
Adaptation to a raptorial mode of life has so profoundly
modified the skeleton of the Falconiformes that much of the
evidence conceruing the origin of the group has been defaced or
obscured.
The most aberrant members of the group are the Catharte.
So markedly do these differ from the other Accipitrine forms that
authorities of no less weight than Garrod and Forbes, for instance,
regarded them as an Order apart therefrom. Thus Garrod placed
them with the Ciconiz and Steganopodes, and Forbes with the
Ciconiz and Tubinares. This was based on a study of the anatomy
of the soft parts—plantar tendons, thigh- and wing-muscles, and
trachea, and on the vestigial condition of the ceca, and in all these
particulars the group is undoubtedly Stork-like.
Mr. Beddard, however, has recently ' expressed his belief in
the derivation of the Falconiformes from a Gruine stem, and has
furthermore brought forward some convincing evidence in support
of his views. The Stork-lke characters of the Falconiformes are
possibly to be traced from their origin low down on the Gruine
stem before the characters common to the diverging branches of
Storks and Cranes began to undergo transformation.
Osteologically, the Falconiformes are certainly more Gruine
than Ciconiine, and here the character most to be depended upon
is found in the skull. It is a comparatively small point, at first
sight, yet it explains the apparently wide differences which
separate the skull of the Cathartz, not only from that of all the
other Falconiformes, but from that of all other members of the
Class Aves.
The Catharte, it will be remembered, have a desmognathous
palate of a quite peculiar type, the maxillo-palatine processes
1 Structure and Classification of Birds. 1898.
312 MR. W. P. PYCRAFT ON THE [ Apr. 15,
being much reduced and widely separated, but, nevertheless,
bridging the palate; and this by means of a pair of strap-shaped —
processes, arising from their inner dorsal borders, and extending
mesially to meet a horizontally expanded plate developed by the
inferior border of the ossified hinder end of the nasal septum.
These processes—the anterior septo-maxillary spurs of Parker—
feebly developed in Rhinochetus and T'etrapteryx, are well seen in
Psophia (P\. XX XIII. fig. 8); and Mr. Beddard was, I believe,
the first to point out their homologies and the part they play in
the formation of the peculiar palate of the Cathartze. Thus, in
describing the palate of Psophia he says, “If these processes were
to be increased in size and to meet a bony internasal septum, we
should have the ‘desmognathous’ skull of the American Vultures.”
It is possible, that at the time Mr. Beddard did not quite grasp
how nearly the Psophiine skull approached the realization of the
modifications peculiar to the Cathartz, since I gather—from his
silence on the point—that there was no trace of an ossified nasal
septum in the skull or skulls which he examined. In all the
skeletons at the Natural History Museum, save one, there is no
septum, which has apparently been lost in maceration. In this
one, it takes the form of a greatly fenestrated plate tapering
forwards to a point, and terminating at the distal fourth of the
external nares. The fenestration in this species is so extensive
that only the hinder end and dorsal border are left, but the hinder
end dips downwards so as nearly to touch the maxillo-palatine
processes. If this septum developed a horizontal plate, as in
the Cathartz, we should have the same type of desmognathism
which is now peculiar to the last-named group.
The nasal septum of the Catharte is more reduced anteriorly
than in Psophia, never extending forwards beyond the posterior
third of the external narial aperture (in the skeleton), and in some
genera is not even visible on a side view of the skull. In
Pseudogryphus and Catharistes it may be studied to best advan-
tage. In the former it extends nearly as far as the middle of the
narial aperture, and is fenestrated much as in Psophia. The bony
tissue which in Pseudogryphus forms the anterior border of the
fenestra is wanting in Catharistes, so that the septum is invisible
when the skull is seen from the side, but, when viewed from
below, the sutures between the horizontal plate of the nasal
septum and the septo-maxillary spurs are plainly visible.
Mr. Beddard, in pointing out the nature of the Cathartine
palate, and the probable source from which it was derived, has
given us the key to a very important problem—the origin of
the Falconiform stem. The Cathartze are the least specialized
members of the group, and, it is interesting to note, are also New
World forms like Psophia. But the low generalized position of
the Cathartz is shown as well by other portions of the skeleton as
by the skull. We may, I think, safely regard the Catharte as
the most primitive of the Falconiformes.
The importance of a correct understanding of the evolution of
1902. | OSTEOLOGY OF THE FALCONIFORMES. 313
the Falconiform palate must be our justification for pursuing this
matter a little further.
The palate of the Cathartz is undoubtedly of an extremely
specialized type, but traces of a like modification are not wanting
in a direction where hitherto they have not been looked for—to
wit, in the Accipitres proper.
The palate of Hlanus, as we have already described, is schizo-
gnathous, a fact first pointed out by Shufeldt ; but the schizo-
gnathism is of a specialized character, being due to the extreme
reduction of the maxillo-palatine processes. If the nasal septum
in the skull of Hlanus be examined, it will first of all be remarked
that it is more complete than in the Catharte, and next that, near
its posterior inferior angle, it gives off a pair of small horizontal
processes, resembling those of Catharte, but relatively smaller ;
these almost touch the maxillo-palatines. A little increase in
the size of these spurs, and the reduction of the anterior portion
of the septum would give us the Cathartine palate. Thus, then,
the palate of Hlanus must be held to represent the high-water
mark of specialization in the direction of schizognathism in the
Accipitres,
It is probable that the palate of Circaétus—or rather the
maxillo-palatines—represents the intermediate type from which
the extremes of schizog- and desmognathism in the group have
been derived. Furthermore, it may profitably be compared with
the paleognathine maxillo-palatine of, say, the Tinamous. It
must be remembered that the great feature of the paleognathine
palate is the enormous size of the maxillo-palatines. In the highly
specialized Zinamus this is much reduced, and in general shape
is not unlike that of Circaétus, but lacks its vertical plate of
spongy tissue. It is important that the intermediate character
of the palate of Circaétus should be recognized, otherwise we
commit ourselves to the admission that the desmognathous
palate of forms like the Falconide or Aquila, for example, were
developed by the resuscitation of an almost defunct organ. In
Circaétus, then, the maxillo-palatines are represented by a pair of
vertical plates of spongy tissue nearly meeting in the mid-ventral
line. They run backwards nearly as far as the antorbital plate
(prefrontal) ; and forwards, then inwards towards the tomium,
so as to leave a palatal vacuity exposing the nasal septum as in
Elanus. Reduction of this type of maxillo-palatine gives us
the highly specialized type of Hlanws, its further increase the
type seen in the Falconide, notably through MWilvago to Polyborus
and Jbycter (Pl. XX XIII. figs. 3, 5), wherein the maxillo-palatines
have attained a relatively enormous size. In the Falconine we
meet with an exactly parallel series, passmg through Harpa to
Herpetotheres (P|. XX XIII. figs. 2-4). In the higher Falcons an
increase in size of the anterior nasal chamber, eventually, in
Falco, developing into a much inflated ossified bulla, has brought
about a considerable reduction in the size of the maxillo-palatine
processes; the fenestrated bulla, seen in such perfection in
Proc. Zoou. Soc.—1902, Vou. I. No, XXI 21
314 MR, W. P. PYCRAFT ON THE [ Apr. 15,
Herpetotheres, ave now veduced to quite vestigial proportions, the
various stages in this reduction being readily traceable through
Cerchneis and Hierofalco (Pl. X XXIII. fig. 1) to Falco.
For further evidence, or rather clues, as to the origin of the
Falconiformes—from an osteological standpoint—one would turn
naturally, to the Serpentariide. In so far asthe skull is concerned,
this would prove disappointing. In its general character it
certainly resembles that of Cariama, but in details it is thoroughly
Accipitrine. The ventral view of the palate resembles that of
Aquila; whilst in the form of the maxillo-palatine processes,
especially with regard to the great size of the antrum of Highmore,
it approaches Leucopternis, Antenor, and Parabuteo. The palate
is indirectly desmognathous. The maxillo-palatines are separated
only by a mere chink, and the desmognathism is effected by the
ossified nasal septum, fusing with the dorsal aspect of the maxillo-
palatines. In the presence of functional basipterygoid processes
it differs from all the Faleoniformes save the Cathartide. In
the trunk skeleton we have already pointed out many Ciconiine
characters, which are not difficult to account for if, as Beddard
and others hold, the Ciconiide and Gruide may claim a com-
munity of descent.
It is certain that the evidence of the skeleton, supported by
the facts which have come to light concerning the myology and
other soft parts, demands that the Cathartide and Serpentariide
must be included with the Falconiformes, though representing,
each, a distinct sub-order. With the Striges the reverse is the
case. Strikingly similar as is the skeleton of the Owls, in many
characters, to that of the Accipitres, it is nevertheless certain
that the nearest allies of this group must be sought among the
“ Picarian” birds. On osteological evidence alone, however, it is
doubtful whether the Striges would ever have been separated
from the Accipitres. The anatomy of the soft parts, however,
seems to prove conclusively the justice of this separation. The
osteological resemblances must be regarded as homoplastic—or, as
some would have it, probably, kinetogenetic.
The relation which the Cathartidee, Serpentariide, and Striges,
severally, bear one to another and to the Accipitres proper, having
now been briefly summarized, we may pass on to the discussion
of the inter-relationships of the last-mentioned group.
This exceedingly difficult problem has lately been attacked by
- Dr. Suschkin. The main results of his study have already
appeared’ **, and, from what he has foreshadowed, it is not
too much to say that his complete Monograph will prove one of
the most valuable and complete contributions to the osteology
of Birds ever published.
1 “ Beitrage zur Classification des Tagraubvogel mit zugrundelegung des osteo-
logischen Merkmale.” Zool. Anzeig. xxii. 1899.
2 “Systematische Ergebnisse osteologisches Untersuchungen einiger Tagraub-
vogel.” Zool. Anzeig. Bd. xxiii. 1900.
3 “ Weitere systematische Ergebnisse vergleichendosteologisches Untersuchungen
der Tagraubvogel.” Zool. Anzeig. Bd. xxiii. 1900.
1902. ] OSTEOLOGY OF THE FALCONIFORMES. 315
My study of this group has convinced me of the soundness of
Dr. Suschkin’s conclusions, as published in the papers to which I
have just referred. But whilst his investigations have extended
over a period of about five years, mine have been limited to a few
months, For this reason I withhold for the present the keys to
the genera, such as have been included in my former papers.
I hope to complete these at a later date.
Before the publication of Dr. Suschkin’s papers I had already
arrived at the same conclusions as are therein expressed with
regard to the position of the Falcones and Polybori, and of
Gypatius and Gypohierax. With regard to Pandion, though I
felt certain it had nothing to do with the Striges, I had not yet
discovered any further clue as to its real affinities.
What follows is practically an embodiment of Dr. Suschkin’s
views in toto: where I have had to interpret him, that is to say
where I have endeavoured to express what I believe to have been
his views, I hope I have done him justice.
The sub-order, then, of the Accipitres is divided into two Families,
the Falconide and the Buteonide.
The family Faleconide is to be divided into two sub-families :
(1) the Falconine, (2) the Polyborine.
The former includes the genera Harpa, Herpetotheres, Micrastur,
Microhierax, Poliohierax, Tinnunculus, Hypotriorchis, Hierofalco,
and falco.
The sub-family Polyborine embraces the genera Milvago, Senex,
Phalcobenus, and Polyborus.
The family Buteonidee is divided into some eleven or twelve
sub-families, though on this point I am not quite clear, as
Suschkin has not definitely expressed himself on this point. But
he would apparently recognize the following: Hlanine, Pernine,
Milvine, Aquiline, Thrasactine, Vulturine, Circattine, Polyborine,
Circincee, Urubitingine, Buteonine, and Accipitrine.
In the Elanine are included Hlanus and Macherhamphus.
In the Pernine, Pernis, Baza, Hlanoides, Leptodon, and Pandion.
But from views he expressed in conversation, he would, I suspect,
probably make a separate sub-family for Pandion—Pandionine ;
and most, I think, will feel this advisable.
The Milvine include Jflvus, Haliastur, and Haliaétus, with,
apparently, /ctinia, Rostrhamus, and Polhioaétus. Haliaétus, there
can be little doubt, has nothing to do with the Eagles. Polioaétus
Suschkin shows to be undeniably distinct from Pandion. The
plantar tendons, as he proved, by a dissection made in this Museum,
are of the Accipitrine type: the skeleton in no way resembles
that of Pandion; on the contrary, the pelvis and breast-bone, so
characteristic in Pandion, bear a quite extraordinary resemblance
to those of Haliaétus. The outer toe is not more reversible than
in ordinary Accipitres. Haliaétus, Thalassactus, and Polioaétus
might well be made to form a separate sub-family, Haliactine.
The sub-family Aquiline includes Aquila, Uroaétus, Spizactus,
Nisaétus, and Lophoaéus. The Thrasaétine contain Morphnus
and Thrasaétus,
316 MR, W. P. PYCRAFY ON THE [Apr. 15,
This brings us to the Vulturine. By most, this sub-family is
regarded as of more importance than is allowed in the present
scheme: Gadow and Sharpe, for example, according it the
rank of a family. That the Vultures have undergone a con-
siderable amount of specialization there can be no doubt; but
it seems equally certain that they are not far removed from the
Circactine. Suschkin recognizes evidence of two distinct branches
in this family—Gypohierax standing at the base of one, and leading
to Neophron and Gyps; Gypaztus at the base of the other, and
leading to Vultur and Otogyps.
The. Circaétinee include Circaétus, aeration” Sani and
Spilornis. Closely allied, and intermediate between it and the
next sub-family—the Circinee—comes the highly specialized
Polyborine. It seems to me that this sub-family might perhaps
as well be included in the Circine, with which, as Dr. Suschkin
shows, it has many characters in common, and in this I can
confirm him.
The Cireinze embrace Circus, CIS ARID EDES, Urotriorcehis, and
Strigiceps.
The Urubitingine I have added on my own responsibility.
Dr. Suschkin speaks of them as isolated forms related to the
Circaétine.
The Buteonine include Buteo, Archibuteo, Tachytriorchis,
Geranoaéius, Rupornis, Leucopternis, and apparently Busarellus,
Butastur, Antenor, and Asturinula.
In the Accipitrine Dr. Suschkin includes Accipiter, Astur,
Melierax, Urospizias, Lophospizias, Scelospizias, and Nisoides.
Though we cannot regard this scheme as final, yet, it must be
admitted, it is one which is in many respects an advance upon
previous arrangements of this most difficult of groups. In its
construction an attempt has been made to follow the lines of
phylogenetic descent, the only satisfactory basis of classification,
yet a peculiarly difficult one in all questions of avian descent,
owing to the lack of annectant fossil forms.
Finally, it is to be noted, the Falconiformes are by no means
so uniformly desmognathous as is generally supposed. Mr,
Beddard has given several exceptional cases, and in the present
paper the list is further extended. But it seems clear that both
desmognathous and schizognathous palates are to be regarded as
modifications of a Gruine-schizognathous type. The Cathartex
have transformed the peculiarities of this type into the unique
desmognathism already described. The forms which still retain
a schizognathous palate have slightly modified the peculiarities of
the original form by loss of the septo-maxillary spurs. Further
specialization has resulted in ee reduction of the maxillo-palatines
to the vanishing point, e.g. Hlanus. But the majority of the
Falconiformes have Peas ‘increased the size of the maxillo-
palatines till they meet in the middle line (a) embracing the nasal
septum between them (indirect desmognathism), or (6) meet
beneath the septum fusing with one another (complete desmo-
1902. ] OSTEOLOGY OF THE FALCONIFORMES, 317
enathism), The various transitional schizognathous types which
occur with considerable frequency, and sporadically, indicate the
steps by which these two forms of desmognathism have been
acquired.
The presence of the hemipterygoid element in all save the
Falcons is a point of considerable interest, serving not only as an
additional index of the high degree of specialization which the
family has undergone, but also to show how a character common
to the members of several widely different orders has been
independently acquired by the modification of a common plan of
structure.
As I have recently pointed out, the Neognathine (Carinate)
palate has been derived directly from the dromeognathous found
only in the Palzognathe. The movement of the palatines towards
the mid-ventral line, whereby they come to underlie the distal
ends of the pterygoids, has caused the latter to segment and the
formation of a pseudo palato-pterygoid joint. The segmentation
of the pterygoids and the fusion of the distal segments thereof
with the palatines relieved them from their function of support-
ing the vomer and threw the work upon the palatines. As a
consequence atrophy of the hemipterygoid ensued; indeed, in
all the members of some sub-orders, e.g. Anseres, Galli, and
in the Falcones, it has been entirely suppressed. When present,
save in a few exceptional cases, it has lost all actual connection
with the vomer, which is borne entirely by the palatines.
The evolution of the Neognathine palate has effected the
following changes :—
1. A shortening of the pterygoids, by the segmentation of their
distal ends and the fusion thereof with the palatines.
2. A lengthening of the palatines, by their forward growth
beneath the maxillo-palatine processes, with which they
originally united, to effect a union with the premaxilla ;
and a change in their position by the movement inwards
to meet in the mid-ventral line.
3. A reduction in the size of the vomer, resulting frequently in
its complete suppression.
But besides changes of position in their relative lengths, the
pterygoids have also undergone a change of function, since
these bones now serve as mere backward extensions of the
palatines, their original function, the support of the vomer,
being transferred as aforesaid to the palatines.
Similarly, the palatines have assumed new functions, in
addition to the support of the vomer. They occupy,
functionally, with the now subordinated pterygoids, the
place of the conspicuous submedian vomero-pterygoid bar
of the Paleognathine skull, wherein, it will be remembered,
they formed but little more than an appendix to the
pterygoid,
318 MR. W. P. PYCRAFT ON THE [Apr. 15,
x. Key To THE OsTEoLOGY OF THE FALCONIFORMES.
A, SKULL.
Upper jaw more or less markedly hooked ; nostrils holorhinal, and when pervious,
associated with functional basipterygoid processes ; temporal fosse, when present,
confined to the lateral aspect of the skull; supra-orbital grooves absent; vomer
blade-shaped ; palate desmo- or schizognathous; an ossiculum lachrymo-palatinum
is never present.
A, Palate indirectly desmognathous, being bridged by union of the septo-maxillary
spurs with a horizontal plate developed by the ossified nasal septum; olfactory
chamber of great size; lachrymal fused with frontal; nares pervious; basi-
pterygoid processes functional; vomer wanting ............... CATHARTA),
B. Palate indirectly desmognathous; basipterygoid processes functional; mavxillo-
palatines with an enormous antrum of Highmore; lachrymal with a very large
supra-orbital process closely applied throughout its whole length to the frontal,
and with its descending process reaching to the quadrato-jugal bar.
SERPENTARIT.
C. Palate directly or indirectly desmognathous or schizognathous ; nares impervious ;
basipterygoid processes vestigial or wanting..................6. ACCIPITRES,
Key to the Lamilies of the Accipitres.
A. Vomer terminating anteriorly in a more or less conspicuous olive-shaped swelling,
closely applied to the maxillo-palatine processes. Palate directly desmognathous; -
antorbital plate (prefrontal) largely developed and with outer border closely
applied to or fusing with the lachrymal; lachrymal without superciliary
plate; squamosal prominence strongly developed; nostrils with a very small
circular or slit-like aperture; ventral aspect of premaxilla with a median bony
ridge; mandible with a ramal vacuity ..................0... FALCONID.
B. Vomer never expanded anteriorly, and never applied to the under surface of the
maxillo-palatines; palate indirectly desmognathous, or schizognathous ; ant-
orbital plate (prefrontal) generally tongue-shaped, often much reduced, or |
articulating, or even fusing by its free end with the distal extremity of the
lachrymal ; nostrils slit-like or pear-shaped, and fully exposing the nasal
septum; lachrymal generally provided with a superciliary plate; squamosal
prominence not greatly developed ...............:cccceceee BUTEONIDE.
Key to the Sub-families of the Falconide.
A. Supra-orbital process of lachrymal extending backwards beyond middle of orbit ;
temporal fossa heart-shaped.............0..0.:tscerese eee Haleonine.
B. Supra-orbital process of lachrymal not extending as far as the middle of the
orbit ; temporal fossa elliptical .....0......ccccieeteeeeee Polyborine.
B. Srernum AND PECTORAL GIRDLE.
Corpus sterni very large, relatively to the size of the girdle, oblong, and with
relatively small anterior lateral processes which bear facets for the articulation of
the anterior sternal ribs; posterior border entire, notched or fenestrated, but the
posterior lateral processes are never very large; articular surfaces for sternal ribs
extend up to or beyond the middle of the sternal plate; coracoid grooves wide, but
shallow, and bordered above by a well-marked “Jip ” ; coracoids slightly overlapping
or only touching one another, and with a well-marked processus lateralis basalis ;
acrocoracoid very large; furenla U-shaped and very broad, with a feebly developed
hypocleideum.
A. Corpus sterni with metasternum produced into a point, bounded by a pair of
notches; furcula articulating with the antero-inferior angle of keel; acro-
: : : a
coracoid not affording an articular surface for furcula ...... SERPENTARII.
B. Corpus sterni with posterior border with two pairs of notches, the outer pair
sometimes becoming closed to form fenestra; keel very deep, extending back-
wards to extreme end of sternum ; coracoid grooves divided from one another in
the middle by a strong ridge; acrocoracoid without articular facets for fureula.
‘ CATHARTA.
1902. ] OSTEOLOGY OF THE FALCONIFORMES. 319
C. Corpus sterni with posterior border notched, fenestrated, or entire; coracoid
grooves overlapping ; acrocoracoid with well-marked facet for articulation of
FULE WI ae een taney cn casual aieegnn aremn ennai ann ACCIPITRES.
Key to Families of the Accipitres.
A. Spina externa and interna well developed; procoracoid large, articulating with
ClAviCle a ey acs ein ca eae ee Pen Ae arn eens VAL CON TD ADs
B, Spina externa only present ; procoracoid articulating with scapula only and
widely separated from clavicle ........0....ccccc0ccceceeeereeeses ACCIPITRIDE.
GC. Pretyic GIRDLE.
Preacetabular ilium very long and with the external lateral border more or less
markedly concave ; pectineal process wanting ; supra-trochanteric processes generally
prominent and raised high above anti-trochanter; pubis, when present, generally
closely approximated to the ventral border of the ischium.
A. Postacetabular ilium shorter than ischium, but without a notch indicating the
two elements posteriorly ...........0..:c00ceeeeeteeeeeeeeseeeeceeeee, SHRPENTARIT.
B. Postacetabular ilium shorter than ischium, and the limitations of the two
elements indicated posteriorly by a deep notch, the ischium being continued
backwards as a sharp spine along the pubis; pubis projecting far beyond ischium,
CATHART A.
C. Postacetabular ilium shorter than ischium, and much deflected ; ischium never
produced backwards into a spine; pubis often vestigial...... ACCIPITRES.
xi. HXPLANATION OF THE PLATES.
Prare XXXI.
Tentral Aspect of the Skull.
Fig. 1. Skull of Catharistes urubu, showing the type of desmognathism peculiar to
the Catharte, wherein the palate is bridged by the union of the horizontal
plate of the nasal septum with a pair of septo-maxillary spurs. Note also
the presence of basipterygoid processes.
Vig. 2. Skull of Elanus ceruleus. The palate is schizognathous. Herein the
maxillo-palatines have increased in size, whilst the septo-maxillary spurs
have completely disappeared. The nasal septum is more complete than in
Catharistes and may be seen lying in the middle line of the anterior palatal
vacuity. Above the inflated region of the maxillo-palatines it sends down-
wards a feeble pair of spurs which nearly touch the maxillo-palatines. The
basipterygoid processes are represented only by a pair of minute prickles.
Tig. 3. Skull of Falco minor. The palate is completely desmognathous. Note the
peculiar form of the vomer, and its relation to the maxillo-palatines. ‘The
antrum of the maxillo-palatines is reduced to the merest vestige.
Fig. 4, Skull of Circaétus gallicus. The palate is schizognathous, and an exaggeration
of that seen in Hlanus. Fusion of the approximated maxillo-palatines
and a downgrowth of and addition to the substance of the nasal septum
would give the palate of Falco. Note the palatal aperture of the antrum.
Fig. 5. Skull of Pseudogyps bengalensis. The palate is indirectly desmognathous.
The palate is bridged by the fusion of the greatly swollen nasal septum
with the widely separated maxillo-palatines.
Fig. 6. Skull of Gypaétus barbatus. The palate is indirectly desmognathous, and
of the same type as in Gyps, but from its less specialized condition shows
how the palate of Gyps has been derived. The vomer is present, the
maxillo-palatines are of great length and widely separated, whilst the nasal
septum can be traced throughout its entire length.
Fig. 7. Skull of Serpentarius serpentarius. The palate is schizognathous, since
the maxillo-palatines, though closely approximated, do not fuse. Fusion
would produce the completely desmognathous type of Falco. The antrum
of the maxillo-palatines is of great size, and the maxillo-palatine processes
are large.
Explanation of letters.
A.p.v. = anterior palatal vacuity. p.a.a. = palatal aperture of antrum.
bp.p. = basipterygoid process. pt. = pterygoid.
map. = maxillo-palatine process. s. = spur of nasal septum.
2.8. = nasal septum. vo. = vomer,
p.a. = palatine.
320
bo
ig. 9.
ie.9)
«
ON THE OSTEOLOGY OF THE FALCONIFORMES. [ Apr. 15.
PLratEe XXXII.
Dorsal aspect of skull of Catharistes wrubu, showing the absence of orbital
processes to the lachrymals and the imperfect nasal hinge. _
Dorsal aspect of skull of Serpentarius serpentarius, showing the large
orbital processes of the lachrymals and their relation to the frontals.
. Dorsal aspect of the skull of Buteo jakal, showing the large, outstanding
orbital processes and the superciliary plate.
Dorsal aspect of the skull of Falco minor, showing the large outstanding
orbital processes of the lachrymal and the absence of a superciliary plate.
Dorsal aspect of the skull of Pandion haliaétus, showing the absence of
orbital processes to the lachrymal. Compare this fig. with the skull of
Catharistes, fig. 1, and note that in Pandion the lachrymal appears on the
surface of the skull.
. Dorsal aspect of the.skull of Microhierax, to show the nasal hinge.
. Lateral aspect of the skull of Wicrohierax fringillarius, to show the nasal
hinge.
. Lateral aspect of the skull of Falco minor, to show the relation of the
lachrymal to the antorbital plate (prefrontal) and the shape of the
nostril. E
Lateral aspect of the lachrymo-nasal region of Pandion haliaétus, to show
the fusion of the lachrymal with the antorbital plate.
ig. 10. Lateral aspect of the lachrymo-nasal region of Serpentarius serpentarius,
to show the relation of the lachrymal to the antorbital plate, and the great
size of the antrum of Highmore.
a.p. = antorbital plate.
l. = lachrymal.
n.h. = nasal hinge.
s. = superciliary plate.
o.l. = orbital process of lachrymal.
Pratt XXXIII.
Lateral view of the upper jaw of Hierofalco gyrfaleo, showing the gradual
suppression of the vertical bullate portion of the maxillo-palatine by the
increasing development of the anterior olfactory chamber (vestibulum
externus).
. Lateral view of the upper jaw of Harpa australis, showmg the more
primitive condition of the vertical bullate portion of the maxillo-palatines
in the Falconine.
Lateral view of the upper jaw of Milvago chimachina, showing the more
primitive condition of the bullate portion of the maxillo-palatines in the
Polyborine.
Lateral view of the upper jaw of Herpetotheres cachinnans, showing the
maximum development of the bullate portion of the mavyillo-palatines
reached by the Faleconine.
Lateral view of the upper jaw of Ibycter ater, showing the maximum
development of the bullate portion of the maxillo-palatines reached by the
Falconine.
Dorsal aspect of the palatal bones of a young Pernis apiworus, to show
the hemipterygeids.
Lateral view of the palatal bones of a young Pernis apivorus, to show the
relation of the hemipterygoid to the vomer.
Lateral view of the lachrymo-nasal region of the skull of Psophia, after
removal of the lachrymal and part of nasal, to show the septo-maxillary
spurs and their relation to the maxillo-palatines and nasal septum.
h.pt. = hemipterygoid.
mex.p. = maxillo-palatine.
nS. = nasal septum.
v.é. = vestibulum externus.
vo. = vomer.
Acanthagyna
rosenbergit, 79.
Accipiter, 289, 297, 298,
304, 306, 307, 311, -
316.
Acesina, 139.
Acherontia
atropus, 205.
Acheta
bimaculata, 94.
capensis, 94,
Achorutes
sp., 1.
Acisoma
panorpoides, 70.
Acomys, 151.
Acreea
alicia, 46.
encedon lycia, 40.
natalica dissociata, 46.
onerata, 46. :
oppidia, 49,
oreds, 45, 46.
orina, 49.
orinata, 45, 46, 48, 49,
51.
porrhasia, 48.
serena rougetit, 46.
toruna, 45, 46. |
uvut, 46. |
vinidia, 46.
zetes menippe, 46.
Acrida
acuminata, 95.
Acridium
ruficorne, 99.
succinctum, 99.
tataricwmn, 99.
Acrobates, 14, 28, 25, 29, |
30.
pygmeus, 14.
Acrydium
INDEX.
Actias
luna, 204.
| Aflurus
Sulgens, 238.
Aipiprymnus, 130.
Agrionoptera
lineata, 69.
mdlaccensis, 69.
nicobarica, 69.
sexlineata, 69.
Alces
bedfordie, 109.
machlis, 109.
resupinatus, 109.
Alestes
grandisquaiiis, 265.
hingsleye, 254.
Allabenchelys, gen. noy.,
234.
longicauda, 234, 235,
237.
Alveolina
melo, 233.
Amauris
albimaculatus, 45.
enceladus, 45.
niavius, 45,
Amblypodia, 139,
Amphieschna
ampla, 64, 78.
_ Amphilius
brevis, 268, 271.
platychir, 268.
Aimphistegina
lessont, 238.
Anabas
maculatus, 269.
Anax
guttatus, 64, 78.
Andinomys, gen.
116.
edax, 116, 117.
nov.,
(Catantops) capicola,
99
_ Anodorhynchus
| glaucus, 107.
Proc. Zoou, Soc.— 1902, Vor. I. No. XXII.
Anodorhynebus
hyacinthinus, 167.
Ancedopoda
latipennis, 94,
Anoplocnemis
tristator, 42.
Antenor, 294,
316.
Anthersea
mylitta, 204.
yama-mai, 204.
310, 314,
Autilocapra, 207, 208,
213, 216, 217.
Apbneeus
hollandi, 45, 46, 49,
51.
oreas, 49,
Aprosmictus
cyanopygius, 169.
Apteryx, 285, 290.
Aquila, 280, 281, 294,
296, 300, 305, 309,
310, 313, 314, 315.
audax, 288,
chrysactus, 288, 295.
VUPax, 299.
Ara
ararauwna, 107.
chloroptera, 168,
macao, 168.
maracana, 168.
nilitaris, 168.
severa, 168.
Archibuteo, 316.
Ardeola
grayt, 268.
Arhopala, 139.
Arvicola
amphihius, 104.
destructor, 104.
Aspongopus
lividus, 42.
—, var,, 42.
nigro-violaceus, 42.
29
322
Astacus
brasiliensis, %.
Jluviatilis, 9, 11.
pilimanus, 9.
Astur, 316.
Asturinula, 316.
Atax
ypsilophorus, 141.
Atella
phalantha, 46.
Aterica
galene, 45.
Attacus
atlas, 204.
cynthia, 204.
orizaba, 204.
ricint, 20-4.
Auchenoglanis
hallayi, 234.
pulcher, 207, 271.
punctatus, 267, 271.
ubangensis, 267, 268.
Azanus
natalensis, 46,
Bairdia, 228.
acanthigera, 229, 230.
amygdatloides, 230.
attenuata, 23\).
crosskeiana, 229, 230.
milne-edwardsii,
230.
fenera, 230.
ventricosa, 230.
woodwardiana, 230.
Balistes, 163.
Baoris
tnconspicud, 47.
Barbus
humeralis, 266.
kessleri, 260.
Barilius
fasciolatus, 266.
ubangensis, 266.
Bassaricyon, 127.
Baza, 515. -
Belenois
agylla, 49.
calypso, 49.
—, var,, 46,
dentigera, 49.
formosa, 46, 50.
ianthe, 49.
instabilis, 46.
mesentina, 46.
rafrayt, 45, 46.
severina injida, 46.
solilucis, 44, 46, 49.
Bernicla
brenta, 160.
pie
229,
INDEX.
Biloculina
depressa,
elongata, 252.
oblonga, 232.
Blatta
egyptiaca, 93.
Brachyccelium, 151.
Brachydiplax
maria, 67.
melenops, 67.
pruinosa, 67.
Brachythemis
contaminata, 63, 66.
Bradypus eek Ae
tridactylus, 129.
Brahmatherium, 215,
218.
Brotogerys
pyrrhopterus, 163.
tui, 168.
virescens, 168.
Bryconxthiops
microstoma, 265.
Bubo, 281, 283.
Busarellus,
316.
Butastur, 316.
Buteo, 280, 281,
316.
jokal, 320.
Buthus
eneas, 223.
bicolor, 223
aint
OHM.
232.
Cacatua
ducorpst, 167.
galerita, 167. :
gymnopis, 167, 170.
hematuropygia, 167.
leadbeateri, 167.
moluccensis, 167.
voseicapilla, 167.
sanguinea, 167.
sulphurea, 167.
triton, 167.
Cacoscelis
ceruleipennis, 190.
compta, 189.
flava, 190.
guianaensis, 189.
marginata, 189.
opacipennis, 190.
testacea, 190.
tibialis, 189.
varipes, 190.
violaceipennis, 191.
Cacyreus
lingeus, 46.
Caica
melanocephala, 168.
294, 10,
289,
Caligula
japonica, 204.
stmla, 204.
Callocvephalon
galeatum, 167.
Calopsittacus
nove-hollandié, 167.
Caloptenopsis
johnstont, 101.
Caloptenus
erassus, 99.
ilepidus, 99.
pinguis, 99.
Calothemis
biappendiculatus, 64,
70.
bivittata, 69.
Calyptorhynchus
naso, 167.
Camacinia °
gigantea, 63, 69.
Cambarus
fallax, 9.
propinguus, 9.
Camelopardalis
giraffa, 52.
Cardium
edule, 142, 148, 155.
Cariama, 290, 292, 294,
Carpenteria
balaniformis, 233.
Catantops
capicola, 99.
humeratis, 99.
Catharistes, 291,301, 508,
5312
Ola.
urubu, 319, 820.
Cathartes, 284, 290, 291,
295, 296; 301, 302,
305, 310.
atrata, 241.
aured, 808.
Catopsilia
jlorella, 46.
Catuna
crithea, 46.
Celxnorrhinus
opalinus, 45, 47.
Centetes, 129.
Cephalophus
sylvicultria, 1.
Ceratomia
amyntor, 205.
Ceratophrys
ornata, 208.
Cerchneis, 314.
Cercopithecus
leucampyx, 237, 238.
otoleucus, 287.
stuhimanni, 288.
Cervicapra
bohor, 138.
Cherocampa
alecto, 205.
elpenor, 205.
Charadrius
Sulous, 262.
Charaxes
bipunctatus, 49.
jasius, 208.
numenes, 45.
tiridates, 45.
Chelzethiops
elongatus, 266.
Cheraps
preissti, 9.
Chilochromis, gen. nov.,
256.
duponti, 234, 236, 237.
Chimachima, 294.
Chinchillula, 116, 117.
sahame, 117.
Chleebora
kelleri, 97.
thalassina, 96.
Chloropza
Jucretiu, +.
Cheeropsis
libericnsis, 239.
Chrysotis
estiva, 168.
agilis, 168.
albifrons, 168.
amazonica, 168.
augusta, 168, 170.
auripalliata, 168.
autumnalis, 16S.
bouquett, 168, 170.
festiva, 168.
guildingi, 168, 170.
qornata, 168.
leucocephala, 168.
levaillanti, 168.
ochrocephala, 168.
salvini, 168.
ventralis, 168.
versicolor, 170.
viridigena, 168.
Circaétus, 288, 294, 305,
313, 316.
gallicus, 319.
Circus, 283, 286, 316.
Clariallabes
gnelas, 235, 237.
Clarias
angolensis, 266.
bythipogon, 266.
Climacobasis, gen, nov.,
85.
lugens, 54, 85, 92.
INDEX,
Coassus
rufus, 217.
Cobus
leucotis, 138.
maria, 138.
Cogia
breviceps, 54-62.
macleayt, 56.
Colias
electo edusa, 46.
Colobus
angolensis, 119.
palliatus, 118, 119.
sharpei, 118, 119.
Colymbus
arcticus, 169.
Conepatus, 111.
Connocheetes
git, Ol.
Conuropsis
carolinensis, 168.
Conurus
acuticaudatus, 168.
eruginosus, 168.
auricapillus, 168.
cactorwin, 168.
holochlorus, 168.
nanday, 168.
ocularis, 168.:
rubrolarvatius; 168.
Coracopsis
nigra, 168.
vasa, 168.
Cratilla
metallica, bo, 68.
Crenis -
boisduvalti, 45.
oceidentaliunr, 45.
Crepidodera
consularis, 205.
flavomaculata, 201.
mayistralis, 202.
(Chalcoides) erichsoni,
202.
Cricetus, idl.
Crocothemis
servidia, 67.
Cryptacrus
comes, 42.
= Vee, 3,
Cryptomima, gen. nov.,
50.
hampsoni, 50, 51.
Curtilla
africana, 94.
Cyanolyseus
patagonus, 168.
Cyanorhamphus
nove-cealandie, \69.
winicolor, 169.
|
Cyclopelta
tristis, 42.
Cymbalopora
poeyt, 233.
tabelleformis, 233.
(Tretomphalus) dz/-
loides, 233.
Cynandra
opis, 45.
Cynictis
levaillanti, 130.
Cynopterus
marginatus, 38.
sphine, 38.
Cyrtacanthacris
pallidicornis, 99.
Cythere
cancellata, 230.
cristatella, 250.
dictyon, 229, 230.
obtusalata, 230.
prava, 229, 230.
rostromarginata, 250.
scintillulata, 230.
stimpsoni, 229, 230.
subrufa, 230.
wyville-thoimsoni, 250.
Cytherella
senitalis, 230.
vesiculosa, 250, 251.
Cytherideis
andrewsi, 229,
Cytheropteron
231.
longicaudatum, 22),
Dail
Damaliseus
tiang, 158.
Darasana, 159.
Dasypus, 227.
minutus, 277.
villosus, 128, 129, 155.
Deilephila
cuphorbie, 205.-
Demodocus, 99.
Deropeltis
erythrocephala, 95.
melanophila, 93.
Devadetta
argyroides, 84,
Dicrocerus
dicranocerus, 217.
elegans, 217,
furcatus, 217.
Dietyophorus
anchiete, 98.
Didelphys
nudicaudata, 180,
Diestogyna
sp., 46.
Diestoeyna
amaranta, 45, 46, 47,
ol.
felicia, 46, 48.
harschi, 48.
Dinidor
tristis, 42.
Dinoceras, 215, 220.
Dinornis, 285.
Diphaulaca
costatipennis, 171.
flavipes, 172, 173.
fruhstorferi, 173.
harold, 173.
pallipes, 172.
Diplacodes
nebulosa, TO.
trivialis, 66.
Dirphia
targquinia, 204, 205.
Discorbina
globularis, 233.
polystomelloides, 253.
Disonycha
angulato-fasciata, 188,
204.
austriaca, 189, 195.
brevicollis, 188.
decemmaculata, 187.
elongata, 187, 197.
Distichodus
fasciolatus, 266.
sexfasciatus, 266.
Distomum, 151.
duplicatum, 141.
margaritarun, 141.
Dolichotis, 131.
patagonica, 277.
Dyspheea
dimidiata, 88.
limbata, 84, 88.
Facles
imperialis, 204.
Echo
modestus, 84, 85,
tricolor, 85,
uniformis, 85.
Welectus
cardinalis, 169.
pectoralis, 169, 170.
westermanni, 169, 170,
171.
Elanoides, 288, 294,
298, 306, 315.
92)
296,
Elanus, 288, 289, 290,
300, 804, 306, 307,
310, 311, 3138, 315,
316.
ceruleus, 319,
INDEX,
Eligmodontia, 116,
Enyalhopsis
petersit, 95.
Kos
riciniata, 167.
rubra, 167.
wallacti, \67.
Equus
burchelli, 32, 37, 38.
prjevalskii, 137.
quagga, 32-58.
Wretis
perpaupera, 47.
Ergolis
enotrea, 46.
Krinaceus, 129.
Hronia
dilatata, 47.
Kuchilichthys
royauxt, 268.
Hupheea
impar, 84, 87, 88.
ochracea, 64, 84, 87.
Hupheedra
eleus, var., 45.
inanuim, var., 45.
spatiosa, var., 45.
vypetina, 45.
Huphysetes, 62.
Huryphene
abesa, 40.
Hurypbymus
crassus, 99.
Eurytela
hyarba, 46,
Eutropius
congolensis, 266.
Falco, 280, 286, 294, 298,
306, 314, 315,
minor, 319, 320.
Felis
concolor pearsoni, 274,
275.
— puma, 218, QA,
275.
onca, 274.
uncia, 137.
Fregata, 303.
Fuligula
ferina, 160.
marila, 158.
Funisciurus
annulatus, 121.
cepapt, 120, 121,
yulei, 120, 121.
Galago
garnetti, 133,
Galinago
celestis, 262.
stenura, 262.
Gastrimargus
determinatus, 97.
marmoratus, 97.
Gaudryina
baccata, 232, 233.
Geranoaétus, 316.
Geranospizias, 516.
Gerbillus, 131.
Globigerina
equilateralis, 233.
bulloides triloba, 233.
dutertret, 233.
helicina, 235.
Glossopsitiacus
concinnus, 167.
Gnathonemus
elephas, 265.
moorit, 265.
rhynchophorus, 265.
Gomphidia
perakensis, 80, 81, 92.
T-nigrum, 81, 82.
Gomphocerus, 96.
Gomphus
consobrinius,
92:
Gonomita
postica, 205.
Gryllotalpa
africana, 94.
Gryllus
bimaculatus, 94.
marmoratus, yar., 97.
Gymnorhina
tibicen, 54.
Gynacantha
plagiata, 79.
rosenhergti, 79.
79, 80,
Gypaétus, 281, 283, 288,
260, 295, 300, 310,
SD wolGs
barbatus, 319.
Gypagus, 280, 291, S01,
306.
Gypohierax, 279, 280,
281, 284, 290, 294,
310, 515, 316.
Gyps, 285, 316.
Gypsina
globulus, 233.
inherens, 232, 238.
Haddonia
minor, 231, 230.
Haliaétus, 280, 281, 294,
300, 304, 305, 310
311, 315.
Haliastur, 289, 294, 315,
Hapalemur
griseus, 128, 135, 185,
simus, 128.
Haplochilus
singa, 269.
Harma
aramis, 45.
capella, 47.
canis, 45.
herminia, 47.
hobarti, 45.
Johnstoni, 45, 47, 51.
lurida, 47.
theobene, 45.
Harpa, 513, 315,
australis, 320.
Hauerina
compressa, 230.
Helictis, 111, 113.
Helladotherium, 215.
Helogale
atkinsoni, 119.
parvula, 120.
undulata, 120.
varia, 119, 120.
victorina, 120.
Helotarsus, 310, 316.
Hemichromis
fasciatus, 270,
Herpestes
parvulus, 120,
pulverulenta, 120.
undulatus, 120.
Herpetotheres, 292, 294,
296, 806, 308, 313,
314, 315.
cachinnans, 320.
Heteracris
bettant, 100.
speciosa, 100.
Heterogamia, 95.
Heterostegina
depressa, 238,
Hetrodes
netersil, 90.
Mierofalco, 289, 314, 515.
gyrfalco, 320.
Hippopotamus
amphibius, 238.
minutus, 238.
Hippopus
hates. 142, 147,161,
162.
Hippotragus
leucopheus, 138.
Huhua
orientalis, 298.
Humbe
tenwicornis, 96.
Hydaspitherium, 215.
INDEX,
Hydrobasileus
extraneus, 65.
Hydrocyon
lineatus, 265.
Hydrophasianus
chirurgus, 51, 261.
Hydropotes
inermis, 217.
Hypochera
zo, 204.
Hypolimnas
misippus, 45,
salmacis, var., 45.
Hypotriorchis, 315.
Hyrax, 129, 133, 134, 135,
136.
919
Ibycter, 515.
ater, 320.
Ictinia, 315.
Ietinus
melanops, 79.
Idionyx
dohrnt, 16, 78, 92.
optata, 78.
yolanda, 76.
| Ischnoptera
melanophila, 93,
Tsotoma
palustris, 13.
| Jagoria
peciloptera, 79.
Kallima
rumia, 46.
| Labeo
falcifer, 236.
greenti, 235, 256, 266.
lukule, 234, 235, 286,
237.
macrostomus, 285, 236,
nasus, 235, 236.
parvus, 235, 236, 266,
Lactica
abdominalis, 174.
apicipes, 178.
bahiaensis, 185.
batesi, 177.
bicolorata, 177, 204.
bilineata, 176.
bogotana, 183.
boliviana, 174.
brevicollis, 182.
capitata, 181.
carinata, 184.
citrina, 180.
clarki, 182.
Co
i)
SU
Lactica
costatipennis, 175,
dichroa, 180.
elegantula, 174.
JSemorata, 181.
flavilabris, 180.
Junerea, 175.
gracilicornis, 185.
impressicollis, 184.
limbatipennis, 177.
lobata, 174.
nigricornis, 185.
paraguayensis, 178.
rufo-basalis, 181.
semifuiva, 180.
seminigra, 176, 204.
strigatipes, 179.
tibialis, 179, 183.
weisei, 179,
Lactina
chalcoptera, 187.
glabrata, 186.
levicollis, 185.
semirugosa, 186.
Lama
huanachus, 2795.
Larus
argentatus, 160.
Lathrecista,
simulans, 69.
terminalis, 69.
Latrodectus
apicalis, 258, 256.
argus, 254, 256.
carolinuin, 253, 256.
cinctus, 255, 256, 257.
conglobatus, 254, 256.
curagaviensis, 251, 254,
257, 261.
curassavicum, 257.
distinctus, 257.
dotatus, 253, 257.
elegans, 257.
ercbus, 254, 257, 258.
JSormidabilis, 258, 257.
geographicus, 251, 254,
257, 260.
geometricus, 248,
252, 256, 257,
261.
hasseltii, 249, 252, 254,
205, 92569) 251 2085
259, 261.
hispida, 254, 258.
hystrix, 251, 262, 253,
258, 261.
indicus, 255, 258.
intersector, 253, 258.
katipo, 252, 255, 256,
258, 261.
lineatum, 253, 258,
205,
326
Latrodectus
lugubre, 254.
lugubris, 254, 258.
mactans, 251, 253, 254,
256, 257, 260, 261.
malmignathus, 248,254,
258.
—, var. tropica, 253.
martius, 254, 258.
menavodi, 252,
256, 259, 261.
nobilis, 257.
oculatus, 254, 259.
ornatus, 259.
pallidus, 251, 253, 259,
260, 261.
perfidus, 259.
quinqueguitatus, 259,
scelio, 255, 258, 259,
261.
schuchii, 259.
spunipes, 260.
thoracicus, 253, 260.
tredecin-guitatus, 247,
251, 252, 253, 254,
255, 256, 257, 258,
259, 260, 261.
tropicus, 260.
variegatus, 258, 260.
variolus, 253, 260.
venator, 254.
verecundum, 353, 260.
zickzack, 252, 260.
zorilla, 253, 260.
Leggada
jerdoni, 39.
Lemur
catta, 130.
Lepadogaster
bimaculatus, 102.
microcephalus, 102.
stictopteryx, 102.
Leptodon, 309, 210, 315.
Lepus
sp., 40.
dayanus, 40.
hainanus, 40, 41.
peguensis, 40, 41.
siamensis, 40, 41,
variabilis, 2.
Testes
orientalis, 92.
ridley?, 64, 92.
udeana, 92.
Leuceronia
argia idotea, 46.
pharis, 46.
255,
Leucithodendrium, 151,
155.
somateri@, 142, 1438,
151, 157, 158.
INDEX,
Leucopternis, 514, 516.
Levinsenia
pygmeun, 158.
Limenitis
camilla, 205.
disippus, 205.
ursula, 205.
Lophaétus, 294, 305,
old.
Lophospizias, 316.
Loriculus
galgulus, 169.
vernalis, 169.
Lorius
domicella, 167.
Loxoconcha
alata, 229, 231.
anomala, 229, 251.
avellana, 231.
honolulicnsis, 229, 231.
Lygzeus
tristator, 42.
Lyriothemis
priapea, 68.
Machxrhamphus, 315,
Macromia
borneensis, 78.
cimcta, 78.
Sumata, 78.
gerstacckert, 76, 77.
westwoodi, 76, 78.
Macropus, 129.
Macroscelides, 225, 227,
228.
proboscideus, 224, 227,
228.
rupestris, 228.
Mahathala, 139.
Manis, 129.
Mantis
undata, 9A.
Marcusenius
pulverulentus, 265.
sphecodes, 254.
Margaritifera,
margaritifera, 141,142,
162, 166.
— mnazatlanica,
162.
maxima, 142, 146, 162.
vulgaris, 141, 147, 162.
Martes, 110.
pentelici, 110.
Mecopoda
latipennis, 94.
Megalopalpus
gymna, 46.
Meles, 110, 111, 113.
paleatiicus, 110,
141,
Meles
(Mustela) paleattica,
110.
Melierax, 297, 316.
Melinda
mercedonia, 45.
Mellivora
ratel, 113.
Melopsittacus
undulatus, 169.
Mephitis, 111, 113.
Mesambria, 98.
Micralestes
altus, 265.
humilis, 265.
stormst, 265, 271.
Micrastur, 3193.
Microdiplax
delicatula, 67.
vittata, 67.
Microgomphus
chelifer, 79.
quadratus, 82.
thoracicus, 79.
Microhierax, 287, 288,
297, 298, 299, 300,
315, 320.
Jringillarius, 320.
Micromerus
affine, 84, 90, 92.
aurantiacus, 84,
hyalinus, 84.
lineatus, 84.
marting, 91.
senuopacus, 91.
signatus, 84.
stigmatizans, 84.
Micromeryx, 217.
Microtus, 131..
sp., LOG.
arvalis, 106, 107.
gregalis, 106, 107.
qivalis, 106, 107.
nivaloides, 106,
(Pitymys) sp., 106.
Miliolina
dicornis, 233.
boueana, 232.
circularis, 232.
— sublineata, 232.
cuvieriand, 232.
ferussaci, 233.
funafutiensis, 231, 232.
gracilis, 253.
insignis, 232.
labiosa, 232.
linneana, 231, 232.
oblonga, 233.
parkeri, 231, 233.
polygona, 233.
seminulum, 233,
Miliolina
subrotunda, 232,
transversestriata, 233.
tricavinata fergweiir-
tana, 232.
undosa, 231, 2388.
Milvago, 289, 308, 513,
315
chimachima, 320.
Milvus, 280, 315.
ater, 289.
Mimomys, gen. nov., 102.
intermedius, 103, 105,
106, 107.
newtont, 103, 105, 106,
107.
pliocenicus, 103, 105,
106, 107.
Miniopterus, 227.
Miracidium, 155.
Modiola
modiolus, 146, 147.
Monotrichtis
perspicua, 45.
safitza campina, 45.
saussure?, 45.
Mormyrops
deliciosus, 265.
Morphnus, 315.
Mus, 131.
concolor, 39.
jerdoni, 39.
Mustela, 109, 110, 111,
113, 114, 130.
flavigulu, 38, 115.
— typica, 38.
joa, 112, W138.
martes, 110, 112, 118.
paleattica, 109-114.
pennant?, 112, 113.
zibellina, 112, 118.
Mya
arenaria, 153.
Mydaus, 111].
Myomyrus
macrodon, 265.
Myopsittacus
monachus, 168.
Mytilus
edulis, 141, 142, 148,
144, 146, 147, 148,
152, W583, Use, WSs,
158, 159, 160, 161,
162, 165, 166.
magellanicus, 162.
Nannocharax
clongatus, 266.
fasciatus, 266.
tenia, 266.
INDEX,
Nannophya
pygmed, 72.
Narathura, 139.
Nasigona, gen. noy., 203.
pallida, 205, 204.
Nasua
narica, 132.
Naudaurelia
cytheria, 205.
tyrrhea, 205.
wahlbergi, 205.
cambesina, 205.
Neoctodon, gen.
114.
simonsi, 115, 117.
Neophron, 281, 288, 290.
300, 316.
Neotoma
cinerea, 115.
Nephrica
holiviana, 192, 194.
brasiliensis, 194, 196,
204.
claveri, 194, 204.
didyma, 192.
Julvicornis, 193.
imitans, 196.
inclusa, 194, 204.
hirschi, 192.
maculipennis, 192, 204.
nigrofasciata, 197.
paraguayensis, 194.
noy.,
sanguinolenta, 193,
204.
staudingeri, 195, 204.
terminata, 195.
unifasciata, 197, 214.
Nephrops
norvegicus, 2-12.
Neptis
melicerta, 46.
nicomedes, 45, 46.
Nesopithecus, 129.
Nesoxenia
metallica, GS.
Nestor
notabilis, 167, 169.
Neurobasis
chinensis, 64, 83, 84,
86. |
Neurocena
ida, 72, 92.
Neurothemis
disparilis, 66.
jfluctuans, 65. |
Sulvia, 65. |
sophronia, 65. |
stiqgmatizans, 65, 66. |
tullia, 63, 66.
Nilasera, 139.
Nisaétus. 315.
Nisoides, 316.
Nubecularia
bradyi, 232.
Nueula
sp., 158.
Nychitona
medusa alcesta, 46.
— immaculata, 46.
Ocnoscelis
boliviana, 191.
Octodon, 114, 115.
Odontopus
notabilis, 42, 43.
Gidaleus
marmoratus, 97.
verticalis, 97.
CHdemia, 157, 159, 160.
nigra, 158, 166.
Onobippidium, 1.
Onychogomphus
geometricus, 80.
— nigrescens, 80.
Onychothemis
abnormalis, 76.
testacea, 75, 76, 92.
Opisthophthahuus, 222.
Orbitolites
duplex, 253.
marginalis, 233.
Orchithemis
pulcherrima, 70.
Oreocinela
dauma, 238.
Orthetrum
sp., 68.
chrysis, 68.
niceville?, 68.
pruinosum, O4, 68.
sabina, 68.
testaccum, 68.
Oryzomys, 116.
Otogyps. 316.
Oxygona
acutangula, 200.
brasiliensis, 200.
luridulus, 200.
nigricollis, 201.
simplex, 200.
Pachytylus
determinatus, 97.
(Cidaleus) marmoratus,
Wc
Pagurus
deformes, 10.
Paleemonetes, 10.
Palzomeryx, 217.
Palornis
derbianus, 169.
docitis, 169,
328
Paleornis
eupatria, 169.
fasciata, 169.
Jinschi, 169, 170.
magnirostris, 169.
schisticeps, 169.
torquata, 169.
Palamneus, 222.
Panchela, 139,
Pandinus, 222.
Pandion, 279,
284, 288,
294, 295,
300, 304,
309, 315.
carolinensis, 296.
haliaétus, 305, 320.
Pantala
flavescens, 63, 65.
Papilio
ajax, 205.
asterias, 20D.
cresphontes, 205.
demodocus, 47.
lormieri, 45, 47.
machaon, 205.
podalirius, 205,
policenes, 47.
troilus, 205.
zolaicon, 205.
Parabuteo, 294, 304, 505,
310, 314.
Parabuthus, 222.
abyssinicus, 222.
flavidus, 222, 225.
granimanus, 222.
heterurus, 222.
hunteri, 222.
liosona, 222.
neylectus, 228.
planimanus, 225.
villosus, 225.
Parastacus
agassizt, 10.
defossis, 10.
hassleri, 10.
safford?, 10.
varicosus, 10.
Payonina, 228.
flahelliformis, 251, 233.
Pelargopsis
gurial, 238.
Pelmatochromis
polyodon, 237.
teniatus, 237.
Peneroplis
arietinus, 233.
pertusus, 233.
(Monalysidium)
draceus, 238.
(—) sollasi, 233.
281, 283,
290, 292,
296, 298,
306, 308,
eylin-
INDEX.
Pentatoma
nigro-violacea, +2.
Perameles, 130, 227.
nasuta, 227.
Pericnemis
stictica, 64.
Peripatus, 11.
Pernis, 281, 283, 284,
289, 298, 309, 315.
apivorus, 3820.
Petasia
anchiete, 98.
Petaurus, 14, 23, 25, 28,
29, 50.
breviceps, 18, 24.
sciureus, 129, 180, 1381,
153, 1385.
Petrocephalus
simuus, 265.
Petrodromus, 225, 227.
Phaethon, 308.
Phalanger, 14, 21, 22, 25,
24, 25, 27, 30.
maculatus, 13.
Phalangista, 130.
Phalcobzenus, 315.
Phascolarctos, 14.
cinereus, 14.
Phascolomys, 25.
Phlceoba
alternata, 96.
rufescens, 95.
Pholas
candida, \47.
Phractura
ansorgii, 269.
hovei, 269.
lindica,
271.
scaphirhynchura, 269.
Phyllotis, 116, 117.
Phymateus
egrotus, 97.
Physeter, 56, 59, 60,
61.
Pinacopteryx
litiana, 46.
Pinna
euglypta, 162,
nigrind, 142, 162.
virgata, 162.
Pionus
chalcopterus, 168.
maximiliant, 168.
Planispirina
extgua, 233.
Planorbulina
acervalis, 233.
larvata, 233.
Platycercus
barnardi, 169.
268, 269,
Platycercus
browni, 169.
elegans, 169, 170.
extimius, 169.
flaveolus, 169.
mastersianus, 169, 170,
171.
pallidiceps, 169.
semitorquatus, 169.
zonarius, 169.
Platy meris
rhadamanthus, 44.
Platypleura
rutherfordi, 44.
Pnorisa
capensis, 96.
Pecilocera
egrota, 97.
Peeocephalus |
gulielmi, 168.
Polioaétus, 315.
Poliohierax, 297, 315.
Polyboroides, 279,
281, 283, 284, 286,
295, 3804, 307, 309,
310.
Polyborus, 280, 283, 287,
289, 294, 304, 308,
313, 315.
Polyphaga
egyptiaca, 93.
Polypterus
bichir, 122.
congicus, 122, 125,
125.
delhezi, 122.
endlicheri,
125.
lapradii,
124, 125.
ornatipinnis, 122.
palmas, 122, 125.
retropinnis, 122.
senegalus, 122,
125.
weeksi, 122, 124, 12.
Polystomella
erispa, 239.
macella, 233.
subnodosa, 238.
Polytelis
barrabandi, 169.
melanura, 169.
Polytrema
miniaceum alba, 232,
233.
Popa
undata, 94.
Potamarcha
congener, 69.
obscura, 69.
122
amy
124,
122
aay
124,
Potamocheerus
penicillatus, 51.
Potorous, 130.
Precis
boopis, 45.
cebrene, 45.
chorimene, 45.
clelia, 45.
gregorti, 45.
terea, 45.
westermanne, 45.
Pristigaster
cayanus, 271.
dolloi, 271.
Procervulus, 217.
Procyon
lotor, 127.
Promeles, 110, 113.
Protoceras, 215.
Protogoniomorpha
temora, 45.
Protorthemis
metallica, 68.
Psammophis
leightoni, 126.
notostictus, 126.
sibilans, 126.
Psephotus
chrysopterygius, 169,
171.
hematonotus, 169.
multicolor, 169.
Pseudathyma
plutonica, 45, 46,48, 51.
stbyllina, 48.
Pseudochirus, 13, 14, 17,
18, 26, 27, 28, 30.
occidentalis, 13, 18:
Pseudogryphus, 280, 291,
301, 312.
californianus, 302.
Pseudogyps, 295.
bengalensis, 319.
Pseudomacromia
luxuriosa, 72.
Pseudophea
impar, 87.
ochracea, 87.
Pseudoplesiops
nudiceps, 270.
squamiceps, 270, 271.
Psittacula
passerina, 168.
Psittacus
erithacus, 168, 170.
Psitteuteles
euteles, 167.
Psophia, 279, 312, 320.
Pteromys, 131.
Ptistes
erythropterus, 169.
Proc. Zoou. Soc.—1902, Vou. I. No. XXIII.
INDEX.
Ptyelus
flavescens, 44.
Pulvinulina
lateralis, 233.
menardi, 233.
repanda, 233.
Putorius, 130.
Pyrrhulopsis
splendens, 169.
Pyrrhura
leucotis, 168.
Querquedula
crecca, 160.
Rhea, 285.
| Rhinocheetus, 312.
Rhinocypha
biforata, 84, 88.
fenestrella, 64, 8+, 88.
heterostigina, 90.
inas, 84, 88, 92.
karschi, 84, 90.
perforata, 90.
petiolata, 8+.
quadrimaculata, 88.
spuria, 88.
tincta, 90.
whiteheadt, 90.
Rhizomys, 131].
Rhodia
fugax, 204.
Rhynchocyon, 225, 227.
Rhyothemis
curiosa, 65.
fulgens, 69.
phyllis, 65.
Rissa
tridactyla, 160.
Rostratula
capensis, 261.
Rostrhamus, 315.
Rotalia
beccarti, 233.
Rupornis, 316.
Saccostomus, 131.
Sagraina
bifrons, 233.
raphanus, 233.
Samia,
ceanothi, 204.
cecropia, 204.
euryalus, 204.
gloveri, 204.
Samotherium, 215, 219.
boissier?, 215.
_ Sarcorhamphus, 280, 291,
301.
329
Sarcorhamphus
equatorialis, 239, 240.
californianus, 285.
gryphus, 239, 240, 241,
242, 243, 244.
Satadra, 139.
Scelospizias, 316.
Schistocera
peregrina, 99.
Sciuropterus, 131.
Sciurus, 131.
atrodorsalis, 38, 39.
caniceps, 39.
castaneaventris gordont,
gordoni, 39.
Scythrops
nove-hollandia, 245.
Senex, 315.
Sericocoris
Johnstoni, 43.
Serpentarius, 278, 279,
280, 283, 284, 287,
288, 289, 290, :
294, 295, 296, 299,
300, 301, 306, 307,
308, 309, 310.
serpentarius, 297, 319,
320.
Sieboldius
grandis, 64, 80, 82, 92.
japonicus, 83.
Sivatherium, 215,
218, 219.
Smerinthus
ocellatus, 205.
tilie, 205.
Somateria
mollissiema, 157.
Sphinx
ligustri, 205.
pinastrt, 205.
Spilornis, 288, 306, 316.
Spirillina
decorata, 233.
inequalis, 233.
limbata, 233.
tuberculo-limbata, 252,
233
Spiroloculina
grata, 232.
tmpressa, 232.
nitida, 232.
Spizaétus, 294, 295, 31d.
Stenobothrus
capensis, 96.
Stomatorhinus
humilior, 265.
Strigiceps, 316.
Struthio, 279, 285.
Surendra, 139.
23
216,
330
Synodontis
decorus, 268.
greshoffi, 268.
pleurops, 268.
Systena
abbreviata, 199.
brasiliensis, 199.
elarki, 198.
punctatissima, 198.
variabilis, 199.
Tachytriorchis, 316.
Nadorna
cornuta, 160.
Tanygnathus
luzonensis, 169.
Tapes, 147, 155, 156, 160,
161, 166.
decussatus, 148, 153,
154, 155, 156, 160.
Vaphronota
gabunica, 97.
Tarsipes, 13, 25, 31.
Tarsius, 129.
Telea
polyphemus, 204.
promcthea, 204.
Tenodera
capitata, 93.
Terias
boisduvaliana, 46.
brigitta zoe, 46.
punctinotata, 45, 46.
Tetracanthagyna
plagiata, 79.
Tetraceras, 218.
Tetragnatha
zorilla, 257, 260.
Tetrapteryx, 312.
Tetrathemis
hyalina, 70, 71.
pulchra, 71, 92.
Tettigonia
flavescens, 44.
Tetyra
comes, 42.
Textularia
agglutinans, 238.
INDEX.
Textularia _
conica, 233.
gramen, 233.
siphonifera, 233.
Thaduka, 139. -
Thais
polyxena, 205.
Thalassaétus, 315.
Tholymis
tillarga, 63, 65.
Thrasaétus, 305, 309, 310,
315
harpyia, 288.
| Tilapia
Fasciata, 271.
stormsi, 270, 271.
Tinamus, 279, 285, 313.
Tinnunculus, 315,
Tirumala
petiverana, 45.
Totanus
hypoleueus, 262.
Traota, 139.
Trichoglossus
Fforstent, 167.
hematodes, 167.
nove-hollandia, 167.
ornatus, 167.
rubritorques, 167, 169.
Trichosurus, 13, 14, 15,
17, 18, 22, 23, 27,
28, 30.
vulpecula, 13, 16.
Tridacna
gigas, 142, 161, 162.
Trithemis
aurora, 63, 66.
intermedia, 66.
trivialis, 63, 66.
yerburii, 66.
Tyriobapta
torrida, 64, 68.
Uroaétus, 315.
Urospizias, 316.
- Urothemis
vittata, 67.
Urotriorchis, 316.
Urubitinga, 310.
THE END.
Vaginulina
legumen, 233.
Vanessa
antiopa, 205.
Verneuilina
spinulosa, 233.
Vertebralina
striata, 233.
Vespertilio
sphins, 38.
Vestalis
amena, 64, 83, 84, 86,
8
7.
Vison, 118.
Viverra
civetta, 130.
tangalunga, 130.
Vultur, 286, 287, 516.
Xenelaphus
bisulcus, 272, 273.
Xestoleberis
curte, 231.
depressa, 229, 231.
margaritea, 231.
Xiphicera
gibba, 98.
haploseelis, 98.
spinulosa, 98.
Ypthima
albida, 45.
granulosa, 45.
Zeltus
antifaunus, 46.
Zizera
antanossa, 46.
Zygonidia
insignis, 74.
malayand, 73, 74.
Zygonyx
ida, 72.
tris, T4, 75.
Zyxomma
petiolatum, 64.
Printed by TayLor and Francis, Red Lion Court; Fleet Street.
THE ZOOLOGICAL SOCIETY OF LONDON.
Tuts Society was founded in 1826 by Sir Sramvorp Rarrrus,
Mr. J. Sasine, Mr. N. A. Vieors, and other eminent Naturalists,
for the advancement of Zoology and Animal Physiology, and for the
introduction of new and curious subjects of the Animal Kingdom,
and was incorporated by Royal Charter in 1829.
COUNCIL.
HIS GRACE THE DUKE OF BEDFORD, K.G., President.
Grorce A. BoutEenerr, Ksa.,
F.R.S.
Toe Hart or Crawrorp, K.T., |
Batuase
Wim E. pr Winton, Hse.
Herpert Drouce, Ese, F.L.S.
Cartes Drummonn, Esa,
Treasurer.
Srr Josrpn Fayrer, Br., F.R.S.,
Vice-President. ;
Dr. Cuarues H. Garry, LL.D.
Dr. AnsERt GuinrHER, F.R.S., |
Vice-President.
Carr. Tas Marquis or Hamirrton,
M.P.
Pror. Grorce B. Howns, D.S8c.,
LL.D., F.R.S., Vice-President.
Lr.-Cot. L. Howarp Irsy.
Sir Harry Jounston, G.C.M.G.,
K.C.B.
Sir Huex Low, G.CU.M.G.
P. Caatmers Mrrcnent, Kse.,
M.A., D.Sc.
EK. Lorr Purutres, Ese.
Howarp Saunpers, Hse., F.L.S.,
Vice-President.
| Puitre Lurtey Scrarer, Ese.,
M.A., D.Sc., F.R.S., Secretary.
Dr. Davin SHarp, F.R.S.
Oxprietp Tomas, Kse., F.R.S.
Dr. Henry Woopwarp, They,
FE.R.S., Vece-President.
2
The Society consists of Fellows, and Honorary, Foreign, and
Corresponding Members, elected according to the Bye-Laws.
The Gardens in the Regent’s Park are open from Nine o’clock A.M.
till Sunset.
The Offices (3 Hanover Square, W.), where all communications
should be addressed, are open from Ten till Five, except on Satur-
days, when they close at Two o’clock p.m.
The Library (3 Hanover Square), under the superintendence of
Mr. F. H. Warernovss, Librarian, is open from 10 a.m. to 5 P.M,
on Saturdays to2 p.m. It is closed in the month of September.
The Meetings of the Society for General Business are held at the
Office on the Thursday following the third Wednesday in every
month of the year, except in September and October, at Four p.m.
The Meetings for Scientific Business are held at the Office twice
a month on Tuesdays, except in July, August, September, and
October, at half-past Hight o’clock p.m.
The Anniversary Meeting is held on the.29th April, at Four p.m.
TERMS FOR THE ADMISSION OF FELLOWS.
Futtows pay an Admission Fee of £5, and an annual Contri-
bution of £3, due on the Ist of January, and payable in advance,
or a Composition of £30 in lieu thereof; the whole payment,
including the Admission Fee, being £35.
No person can become a Feniow until his Admission Fee and
First Annual Subscription have been paid, or the annual payments
have been compounded for.
Fentows elected after the 30th of September are not liable for
the Subscriptions for the year in which they are elected,
PRIVILEGES OF FELLOWS.
Frettows have Personal Admission to the Gardens with Two
Companions daily, upon signing their names in the book at the
entrance gate.
FELLows receive a Book of Saturday and a Book of Sunday Orders
every year. ‘These Orders admit two persons to the Gardens on each
Saturday and two on each Sunday in the year. But the Saturday
3
Orders are not available if the Frriow shall have used his privilege
of personally introducing two companions on the same day.
Ferttows also receive every year Twenty Free Tickets (Green),
each valid for the admission of one adult any day of the week,
including Sunday. Children’s Tickets (Buff) can be had in lieu of
Green Tickets in the proportion of two Children’s Tickets to one
Adult’s. These Tickets, if not made use of in the year of issue, are
available for following years.
Frttows, if they wish it, can exchange the Book of Saturday
Orders for Twenty Green Tickets available for any day. The Book
of Sunday Orders can also be exchanged for a similar packet of
Twenty Tickets. These bocxs must, however, be returned entire,
and the exchange can only be made during the year of their issue.
The annual supply of Tickets will be sent to each Frttow on the
Ist of January in every year, on his filling up a form of Standing
Order stating in what way they should be made up, and to what
address they should be sent. Forms for this purpose are supplied
on application.
The Wire of a FrLtow can exercise all these privileges in his
absence.
Frttows have the privilege of receiving the Society’s Publications
on payment of the additional Subscription of One Guinea every
year. This Subscription is due upon the Ist of January and must
be paid before the day of the Anniversary Meeting, after which
the privilege lapses. Frxtows are likewise entitled to purchase the
Transactions and other Publications of the Society at 25 per cent.
less than the price charged to the public. A further reduction of
25 per cent. is also made upon all purchases of Publications issued
prior to 1871, if above the value of Five pounds.
Fettows also have the privilege of subscribing to the Annual
Volume of the Zoological Record for a sum of £1, payable on the
1st July in each year, but this privilege is forfeited unless the
subscription be paid before the 1st of December following
They may also obtain a TransreraBLe Ivory Ticker admitting
Two Persons, available throughout the whole period of Fellowship,
4
on payment of Ten Pounds in one sum. A second similar ticket
may be obtained on payment of a further sum of Twenty Pounds.
Any Frriow who intends to be absent from the United Kingdom
during the space of one year or more may, upon giving to the
Secretary notice in writing, have his name placed upon the
“ dormant list,” and will be thereupon exempt from the payment of
his annual contribution during such absence.
Any Frtxow, having paid all fees due to the Society, is at liberty to
withdraw his name upon giving notice in writing to the Secretary.
Persons who wish to become Fellows of the Society are requested
to communicate with the undersigned.
PHILIP LUTLEY SCLATER, M.A., D.8c., F.R.S.,
Secretary.
Hanover Square, London, W.,
August, 1902.
MEETINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON
FOR
SCIENTIFIC BUSINESS.
(AT 3 HANOVER SQUARE, W.)
Session 1902-1903.
1902.
Turspay, Novemprr 4 and 18 | ‘Tvuxspay, Decemprr 2
1903.
Turspay, JANUARY 20 Tunspay, Aprin .. 21
“ Frervary 3 and 17 =46 May .... 12and 26
” MAR CHAS Sons lay. A CUO G5 o IG
The Chai will be taken at half-past Hight o'clock in the Evening
precisely.
LIST OF THE PUBLICATIONS
OF THE
ZOOLOGICAL SOCIETY OF LONDON,
Tue scientific publications of the Zoological Society of London
are of two kinds—“ Proceedings,” published in an octayo
form, and ‘‘ Transactions,” in quarto.
According to the present arrangements, the “ Proceedings”
contain not only notices of all business transacted at the scien-
tific meetings, but also all the papers read at such meetings
and recommended to be published in the “ Proceedings” by
the Committee of Publication. A large number of coloured
plates and engravings are attached to each annual volume of
the “ Proceedings,” to illustrate the new or otherwise remark-
able species of animals describedin them. Amongst such
illustrations, figures of the new or rare species acquired in a
living state for the Society’s Gardens are often given.
The “ Proceedings” for each year are issued in four parts,
on the first of the months of June, August, October, and
April, the part published in April completing the volume
for the last half of the preceding year. Commencing from
January 1901 they form two half-yearly volumes.
The ‘ Transactions” contain such of the more important
communications made to the scientific meetings of the Society
as, on account of the nature of the plates required to illustrate
them, are better adapted for publication in the quarto form.
They are issued at irregular intervals.
Fellows and Corresponding Members, upon payment of
a Subscription of One Guinea before the day of the Anni-
versary Meeting in each year, are entitled to receive all the
Society’s Publications for the year. They are likewise
entitled to purchase the Publications of the Society at 25 per
cent. less than the price charged for them to the Public. A
further reduction of 25 per cent. is made upon purchases of
Publications issued prior to 1871, if they exceed the value of
five pounds.
Fellows also have the privilege of subscribing to the
Annual Volume of the Zoological Record for a sum of £1
(which includes delivery in the United Kingdom only),
payable on the Ist July in each year; but this privilege
is forfeited unless the subscription be paid before the Ist of
December following.
The following is a complete list of the publications of
the Society already issued. They may be obtained at
the Society’s Office (8 Hanover Square, W.), at Messrs.
Longmans’, the Society’s publishers (Paternoster Row, H.C.),
or through any bookseller. —
| August, 1902. |
TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON.
4to. 15 vols. and Index. Prete
Vol I, containing 59 Plates.... (1853-35) .... £3 13
po smells fo iierrmmmarere a (lice se vba (0)
Me UE ee 63, . (1842-49) .... 3 8
Ne ae ba 4° Te ets ee CLIO GO) EG
OE Me G7) nhs Me isepsGaye we hua e ee
i OS) , COD) o.5. 1 F
NU . xe (USES 7) a ong LO
HAVTT Eiht wc BDIse haat Aur ISO ane ao TS
aie oa 4 QO anes. LOO (arene el el
PP ia ar! ROB NE (1877-79) .... 10 0
index, Vols M Xs yesh mein cee SEO) yon Wl) a
Vol. XI, containing 97 Plates.. (1880-85) .... 9 12
cal i pV ies6 200) Matas ee
aI eer Gon EM sor Lon nam ieanG! ie
per OXGLIV A op on (UISDE-DS). 5 5
AEN OMS we Se =1001) een a iG
XG eile oO, es K(Heb lOO): 4 Ons
Y SOUL eB oh OB Ke (ee TOO) os Oni
MeV ae ou Oia ht OceMIGOl ame Oy
PETBXON ALY RCA AS Bo om (Ajmal IS02) 05 © 2
TONGVATE, Que EN MINE. Wk OA UT aeek nhs oh 011
SO, Guy oh ee aCe OO, 0) 1
”)
OWNOODWOHDOWODOAWAWOOWOWOR
Price to the
Public.
ocoocooqoooceoa
oo°occeo
*
* Part V. of Vol. XVI., containing Mr. Mudge’s paper on the Neelees of is Tongue of
Parrots, is not yet ready and will be issued Buea
PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND
CORRESPONDENCE OF THE ZOOLOGICAL
LONDON. 8vo. 2 vols. (Letterpress only).
Part I.
IL.
7
1 vol. 8vo.
”
1830-81,
1882.
es eeceecteose eee
Price to
Fellows.
As. 6d.
SOCIETY OF
Price to the
Public.
6s.
6s.
PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.
8vo. 15 vols. (Letterpress only) and Index.
Price to Price to the
(First Series.)
Price to Price to the
Fellows. Public, Fellows. Public.
Part 1. 1833. l vol. 8vo. 4s. Gd. .. 6s.F | Part 1841.1 vol. 8vo. 4s. 6d. .. 6s.F
» IL. 1884. 45 4s. 6d. .. 6s. <9 X. 1842. 59 4s, 6d. .. Gs.
SLU S3o. > 4s. 6d. .. 6s. e XI. 1848. As. 6d... Os.F
» LV. 1836. 9 4s, 6d. .. 6s. » XID. 1844. As. 6d. .. 6s.
«| No dhe Bis 9 4s. 6d. .. 6s. 5 S45. » 4s. 6d. .. 6s.
wy Walle Ie tak. 4s. 6d. .. 6s. » ALV. 1846. = 4s, Gd. .. 6s.F
oy AUIS Skee) i As, 6d. .. 68.+ XV. 1847. a 4s. 6d. .. 6s.
» VIII. 1840. "9 ASNGG a roan OSs Index 1830-1847. . 4s, 6d. .. 65.
8vo. 13 vols. and Index. (Second Series.)
Letterpress only. With Plates coloured.
Price to Price to the Price to Price to the
Fellows. Public. Fellows. Public.
Part XVI. 1848. 1 vol. 8vo. 4s. 6d. Gsiin Rasienscie 1 0) (8° 5. Sligo
% XVII. 1849. “ 4s. 6d. OS ature carrie Ihe Orgs Le Or
» XVIII. 1850. i‘ 4s, 6d. LOPE Enc 1 BS oe LAST Or
3 XIX. 1851. bs As, Gd. G85 Meas ners ae Os is es) Lo sO
5 XX, 1852. % 4s. 6d. GS a ile aya cieks OW5L59 ebe Oy
XXI. 1853. 5 4s, 6d. SF MME cee ae 0) Tis} 9 @) 1 4 OF
5 XXII. 1854. mn As. 6d. OST ante oper 019 6 letoe Oy
» WXITTL 1855. “ 4s. Gd. Ost Bc SAE cee ei © 118 OF
EOE Jhet op 55 As, 6d. Ges) (FONE LORS De ey;
z XXV. 1857. 1 4s, Gd. GS. a ahecerable LORS 1B ee Gap
» S&XVI. 1858. a, 4s, 6d. Lo Mewar he Gc IPH © Py iy hy
» XXVII. 1859. as As. 6d. Ostia Recents Te JHE. & 2) 2205
XXVIII. 1860. Ms 4s, Gd. Ok ga Gouce: TINE 6 2 2 Of
Index 1848-1860. 7 As. 6d. 6s.
t Ont of print.
PROCEEDINGS OF THE SCIENTIFIC MEETINGS OF THE
ZOOLOGICAL SOCIETY OF LONDON. 8vo. 30 vols. and 3 Indices.
Letterpress only. With Plates uncoloured. With Plates coloured.
Price to Price to the Price to Price to the Price to Price to the
Fellows. Public. Fellows. Public. Fellows. Public.
USGI wAS Oda ees OSE eae: 9s. Pree epi Coke, seh esice Sas. 9d. .... 458.1
SGQs As Od rane OSuie net 9s. een Kees al BES; Be odes 45s.7
USGS 5 5 4 Gro o 866 (OS ooboce 9s, Reraal Peaee ar Or ak, Bs Sb oac5 4K
SCRE eASMGIS asm (OSs. «aie 9s. Ge WD Sar ie ares SB Buh once CESS P
NSGomM ra seiGds ool Ose sek 9s, Reastay Shi hae toten Si Sh ona AS
HSS 56 4's Gh soa0 GSE 555006 9s. oer el iPen aetiseesteteter Ba Bl coos 45s,
RSG fpapenmin re r a rareetie ou aie ent se. 9s. Bere enn Lasoie apeit elec eet 80h bo 5. 4m
SG Shree. Moneere eects cttw tee Qs. Ba lel eee aS SB Wh oco0 Zim
TCKGIS MR, Sissel tices CR) CeO gh Ane ge 9s. Port iben all OU tang Auta BB Bh ono. GOS
SOR eras icn Gere ni blere Kas eunle 9s. Na SAMMI SN ote cana BG Bh son. 2iR,
ide wISGINS(0i ap aan cere AS TOG Oma OS.
1 ph Ne aR Re A ee 9s. RE ai Senne ars Sosy Oden os!
ASAD eae ea Ata te eater caret oeses 9s. LAUR S| SORE Nae oes. 9d. .... 408.1
TGV)" 4h ois Reena Beek et aCe ae 9s, ei rein bese pe ea Oo A des. Od. .... 458.
UCT RUE Ane ORCA Pan rent 9s. SA aod atin ne a0 POLO cone Aki;
UGH AD pee aren iexaeestece ce eae cia eats 9s. Eg eo ART ee open te 36s. Meee ASs
TISYAG? bec Bee Oru Se Taped a Rate err 9s. Fes HILO mune ea eae ak sooo Sek
ANS i ieey ene Maron teda ine eV erey Ones aN 9s. aac hO Se) a cerry 36s, oooo Ake
ESAs! ch eel ee soe Raa Al eS RC LE 9s. mL Sate tt ee 36s. soae Akeis,
TSN AG) A Rar el ce ale i aa et a ey 9s. arte ie Seaceecryenres 36s. sono Eke
SSO Pet a euro Mus mente Eas Qs. Weep ED Gi ok ese 36s. soo 4Seh
IndexsplS Al —ISSOMea ens Ace GS:
Octo 1 ua ac Geen ear mee Mee tare ieall ry aire 9s. eH OSG nase ar 36s. ASS:
TU CkS I) Mac Re teas Ree Ade ese Be aes V 9s. paca Stach 36s. cobe 4keK.
ROSES re soe Rpt Mente 9s. Rea Bal kee) 36s. sooo 4h:
MO SANE Mabe MSE NA etic Nace NL beta 9s. nea Tt Sanaa OA aah 36s. As
ARGS OMe a ne unr wees aero. 9s. Sei eS 36s. ASS:
USS OR reece tae uae arnet am te tn a accent 9s. Sebel KOA ay NN 36s. ALGO
ISS eeer cinemas at Ace a Me RE 9s. SRO OBEY Se 36s. sid .d ALS
TUCKSRS 28 ites Ree BB ne See ae) a aU 9s. pies Pea ie LEDS Bs mabe Wee aay Aan oo Hes
TENSION ao ian atta (Nea an hale a ae 8 9s. eNO SRO EAA 36s, ARs
BS 9 Oey ek connie Potap crane. 9s. Bhs tavnterd ALON Mente te ie 386s. sooo Shes,
Index. SSI 1S90y Foes elu ests clei: 4s. Gd. ..., 6s.
* No perfect copies in stock. t Out of print.
PROCEEDINGS or tHe GENERAL MEETINGS ror SCIENTIFIC
BUSINESS or tuzr ZOOLOGICAL SOCIETY OF LONDON.
8vo. 12 vols.
Price to Price to the
Fellows. Public.
HN ONT eereea eMart Muna artae sted den Mn gleere article oun aapeien a aban act SOS ane 485
SUPSEO POA ah ain. Taco ety ror tl Gale PUI CREIS CChS CDI RE sie caernenin (ah here earer DOS TN og 48s
SO SRM Mei ap ccver emit rattars anencuescrm veerskezoysicestate cassie iniey-ei oe oralte rea ettenav a Sacre BOS ra ae 48s
US OA ee Aree cuaWayet vis delon oacsin ie eriievel eiece ceiaieesle- 6 ace ators Gace aletns mie Row erens SOS5- yes 48s
TEENS SS ace 25 cece La Ete EE Sh Cae RES ear ea Tac er eR SOS aes 48s
ELS QO eveereret vareaer tute netbeans seals de. acata eee: aaa lus atenasaieae ater aver cules Sosne eens 48s
MS Gay rea Une earch KeReaGT Osta se eGeue ss 6 SlenaTerbvosuae WNievuin a badiaar a ieeenauconcusliodeds SOSs Aen A8s
ANS OST repre ema ce aah ciate ovis saves cuoeices altaya: ortaunc cos veeaile a emade onaeare nate, aus BOSS eee 48s
NSO OMA ateerc asian a tare esas Metter ocel sade ola tha level kc scevorelers RI plone A8s,
LD OO eee ee cats mevels\evietian ata craeti ebor er aytauesarees) si's, w. deb et ara weal ae eel Goo SESH awe A8s.
HOO NS yo lege lircataenre ise versa vtscn siovaistedi reser soldi a'si-s asleie) Oie.a/aia-scstare alla ISHS Sloman a> 24s,
yy MUL: AAW ae See Leen nS ines ilmenre aye issn en 2ds.
LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS.
List of the Vertebrated Animals now or lately Living in the Gardens
of the Zoological Society of London. (Highth Edition.) Cloth.
8vo. 1883. Price 3s. 6d.
List of the Vertebrated Animals now or lately Living in the Gardens
of the Zoological Society of London. (Ninth Edition.) 8vo.
1896. Price 4s. 6d.
THE ZOOLOGICAL RECORD.
The Zoological Record for the years 1864-1900. Thirty-seven
volumes. 8vo. Price £15, Net.
The Zoological Record, Volume the Thirty-first. Being Records of
Zoological Literature relating chiefly to the year 1894. Edited
(for the Zoological Society of London) by Davin Suarp, Ksq.,
M.A., F.R.S., F.Z.8. London, 1895. 8vo. Price 10s.
The Zoological Record, Volume the Thirty-second. Being Records of
Zoological Literature relating chiefly to the year 1895. Edited
(for the Zoological Society of London) by Davin Suarp, Esq.,
M.A., F.B.S., F.Z.S. London, 1896. 8vo. Price 10s.
The Zoological Record, Volume the Thirty-third. Being Records of
Zoological Literature relating chiefly to the year 1896. Edited
(for the Zoological Society of London) by Davin Smarr, M.A.,
F.RS., F.Z.8., &. London, 1897. 8vo. Price 30s.
The Zoological Record, Volume the Thirty-fourth. Being Records of
Zoological Literature relating chiefly to the year 1897. Hdited
(for the Zoological Society of London) by Davin Suarp, M.A.,,
F.BS., F.Z.8., &. London, 1898. 8vo. Price 30s.
The Zoological Record, Volume the Thirty-fifth. Being Records of
Zoological Literature relating chiefly to the year 1898. dited
(for the Zoological Society of London) by Davin Suaxp, M.A.,
E.R.S., F.Z.8., &. London, 1899. 8vo. Price 30s.
The Zoological Record, Volume the Thirty-sixth. Being Records of
Zoological Literature relating chiefly to the year 1899. Hdited
(for the Zoological Society of London) by Davin Saarp, M.A.,
F.R.S., F.Z.8., &c. London, 1900. 8vo. Price 30s.
The Zoological Record, Volume the Thirty-seventh. Being Records
of Zoological Literature relating chiefly to the year 1900.
Edited (for the Zoological Society of London) by Davin Suarp,
M.A., F.R.S., F.Z.8., &. London, 1901. 8vo. Price 30s.
Catalogue of the Library of the Zoological Society of London.
(Fourth Edition.) Cloth, 8vo. 1887. Price 4s.
These publications may be obtained at the Sociery’s Orrice
(8 Hanover Square, W.), at Messrs. Lonemans’ (Paternoster How,
E.C.), or through any bookseller.
THE ZOOLOGICAL RECORD.
——-059300——_.
[HE object of the ZootoercaL RucorD is to give, by means of an
annual Volume, complete lists of the Works and Publications
relating to Zoology in all its branches that have appeared during
the year preceding the issue of the Volume, together with full
information as to the points they deal with, arranged in such a
manner as to serve as an Index to the literature of Zoology in all
parts of the globe, and thus to form a repertory which will retain
its value for the Student in future years.
The ‘ Zoological Record’ is published for the Society by Messrs.
Gurney and Jackson at the price of 30s. per volume. But all
Members of the Zoological Society of London have the privilege
of receiving it, including the cost of delivery (within the United
Kingdom), at a subscription price of 20s. per annum. This Sub-
scription is due on the lst of July in every year, and the privilege
of Subscription is forfeited unless the amount be paid before the
1st of December following.
The Zoological Society, having purchased the entire stock of
the ‘ Zoological Record,’ are able to supply complete sets. ‘The
thirty-seven Volumes to the end of the nineteenth century, and the
Index-Volume in addition, will be supplied for £15. Volumes of
any single year (exclusive of the last five volumes and Vol. 6)
can likewise be supplied at 10s. per volume.
Members of the Society wishing to subscribe to the ‘ Record’
are requested to apply at this office for a Form, to be returned
when filled up and signed by the subscriber. In order to facilitate
the payment of the subscription, a Banker’s Order Form is also
furnished to those who prefer that mode of payment. ‘This order,
when filled up and signed, should be sent to the Society’s office for
registration ; 1t will then be sent to the Agents named therein.
Learned Societies and Institutions and members of the former
Zoological Record Association are permitted to subscribe to the
‘Record’ on the same conditions as are accorded to Members of
the Zoological Society.
P. L. SCLATER,
Secretary.
August, 1902.
ZOOLOGICAL SOCIETY or Lonpon,
& Hanover SQuaxn, W
LIST OF VOLUMES or vxe “ZOOLOGICAL RECORD.’
The Record of Zoological Literature, 1864. Volume First.
Edited by Auserr C. L. G. Giwrunr, M.A., M.D., Ph.D., F.Z.8., &e.
London, 1865. Price 10s.
The Record of Zoological Literature, 1865. Volume Second.
Edited by Atperr C. L. G. Gunrnmr, M.A., M.1)., Ph.D., F.Z.8., &e.
London, 1866. Price 10s.
The Record of Zoological Literature, 1866. Volume Third.
Edited by Atperr C. L. G. Ginter, M.A., M.D., Ph.D., F.B.S.,
F.Z.8., &c. London, 1867. Price 10s.
The Record of Zoological Literature, 1867. Volume Fourth.
Edited by Auserr C. L. G. Gunrner, M.A., M.D., Ph.D., F.R.S.,
F.Z.8., &c. London, 1868. Price 10s.
The Record of Zoclogical Literature, 1868. Volume Fifth.
Edited by Auserr 0. L. G. Ginenmr, M.A., M.D.; Ph.D., F.RS.,
F.Z.S., &c. London, 1869. Price 10s.
The Record of Zoological Literature, 1869. Volume Sixth.
Edited by Atpert C. L. G. Ginruyr, M.A., M.D., Ph.D., F.RB.S.,
F.Z.8., &c. London, 1870. Price 30s.
The Zoological Record for 1870; being Volume Seventh of the
Record of Zoological Literature. Hdited by Atrrep Newrov, M.A.,
E.RB.S., F.L.S., V.P.Z.8., &e. London, 1871. Price 10s.
The Zoological Record for 1871; being Volume Fighth of the
Record of Zoological Literature. Kdited by Atrrep Newron, M.A.,
E.R.S., F.L.S., V.P.Z.8., &. London, 1873. Price 10s.
The Zoological Record for 1872; being Volume Ninth of the
Record of Zoological Literature. Edited by Atrrep Newron, M.A,,
E.R.S., F.L.S., V-P.Z.8., &e. London, 1874. Price 10s.
The Zoologieal Record for 1873; being Volume Tenth of the
Record of Zoological Literature. Edited by Epwarp Catpwet
Ryz, F.Z.8. London, 1875. Price 10s.
The Zoological Record for 1874; being Volume Eleventh of
the Record of Zoological Literature. Hdited by Epwarp Catpwett
typ, F.Z.8., M.E.S. London, 1876. Price 10s.
The Zoological Record for 1875; being Volume Twelfth of the
Record of Zoological Literature. Edited by Epwarp Catpweutt Ryz,
F.Z.8., M.E.S. London, 1877. Price 10s.
The Zoological Record for 1876; being Volume Thirteenth of
the Record of Zoological Literature. Edited by Epwarp Catpwern
Ryz, F.Z.8., M.E.S. London, 1878. Price 10s.
The Zoological Record for 1877; being Volume Fourteenth of
the Record of Zoological Literature. Edited by Epwarp Catpwett
tye, F.Z.S., M.E.S. London, 1879. Price 10s.
The Zoological Record for 1878; being Volume Fifteenth of
the Record of Zoological Literature. Edited by Epwarp Catpwstt
Rye, F.Z.8., M.E.8. Londou, 1880. Price 10s.
The Zoological Record for 1879; being Volume Sixteenth of
the Record of Zoological Literature. Hdited by Epwarp Catpwett
Ryu, F.Z.8., M.E.8. London, 1881. Price 10s,
The Zoological Record for 1880; being Volume Seventeenth of
the Record of Zoological Literature. Edited by Epwarp Canpwet.
Ry, F.Z.8., M.E.S. London, 1881. Price 10s.
The Zoological Record for 1881; being Volume Eighteenth of
the Record of Zoological Literature. Edited by Epwarp CaLpweEnt
Ryz, F.Z.S., M.E.S. London, 1882. Price 10s.
The Zoological Record for 1882; being Volume Nineteenth of
the Record of Zoological Literature. Edited by Epwarp Catpwent
RYE, F.Z.8., M.E.S. London, 1883. Price 10s.
The Zoological Record for 1883; being Volume Twentieth of
the Record of Zoological Literature. Edited by Epwarp Catpwrzn
Ryg, F.Z.8., M.E.S. London, 1884. Price 10s.
The Zoological Record for 1884; being Volume the Twenty-
first of the Record of Zoological Literature. Edited by F. Jurrrey
Brett, M.A. London, 1885. Price 10s.
The Zoological Record for 1885; being Volume the Twenty-
second of the Record of Zoological Literature. Kdited by F. Jurrrny
Brett, M.A. London, 1886. Price 10s.
The Zoological Record for 1886; being Volume the Twenty-
third of the Record of Zoological Literature. Edited by Franx EK,
Brpparp, M.A., F.Z.8. London, 1887. Price 10s.
The Zoological Record for 1887; being Volume the Twenty-
fourth of the Record of Zoological Literature. Edited by Franx E,
Bepparp, M.A., F.Z.8. London, 1888. Price 10s.
The Zoological Record for 1888; being Volume the Twenty-
fifth of the Record of Zoological Literature. dited by Franx E.
Brepparp, M.A., F.Z.5. London, 1890. Price 10s.
The Zoological Record for 1889; being Volume the Twenty-
sixth of the Record of Zoological Literature. Edited by Franx HK.
Brpparp, M.A., F.Z.S8. London, 1890. Price 10s.
The Zoological Record for 1890; being Volume the ‘Twenty-
seventh of the Record of Zoological Literature. Kdited by Frank
KH. Bepparp, M.A., F.Z.S. London, 1892. Price 10s.
The Zoological Record, Volume the Twenty-eighth; being
Records of Zoological Literature relating chiefly to the year 1891.
By J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger,
W. A. Herdman, B. B. Woodward, C. Warburton, R. I. Pocock,
D. Sharp, E. A. Minchin, A. Willey, and 8. J. Hickson. Hdited by
D. Saarp, M.A., F.R.S., F.Z.8., &. London, 1892. Price 10s.
The Zoological Record, Volume the Twenty-ninth; being
Records of Zoological Literature relating chiefly to the year 1892.
By J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger,
W. A. Herdman, B. B. Woodward, R. I. Pocock, D. Sharp, F. A.
Bather, Florence Buchanan, S.J. Hickson, and R. Hanitsch. Edited
by D. Saanp, M.A., F.R.S., F.Z.S., &. London, 1893, Price 10s.
_ The Zoological Record, Volume the Thirtieth; being Records
of Zoological Literature relating chiefly to the year 1895. By J. A.
Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger, W. A,
Herdman, B. B. Woodward, R. I. Pocock, D. Sharp, F. A. Bather.
Florence Buchanan, 8. J. Hickson, and R,. Hanitsch. LKdited by
D. Saarp, M.A., F.R.S., F.Z.8., &. London, 1894, Price 10s.
The Zoological Record, Volume the Thirty-first ; being
Records of Zoological Literature relating chiefly to the year 1894.
By J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger,
W. A. Herdman, B. B. Woodward, D. Sharp, F. A. Bather, Florence
Buchanan, and R. Hanitsch. Edited (for the Zoological Society of
London) by D. Swarr, M.A., F.R.S., F.Z.8., &c. London, 1895.
Price 10s.
The Zoological Record, Volume the Thirty-second; being
Records of Zoological Literature relating chiefly to the year 1895.
By J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger,
W. A. Herdman, B. B. Woodward, R. I. Pocock, D. Sharp, F. A.
Bather, Florence Buchanan, R. T. Giinther, R. von Lendenfeld,
W. F. Hume, and F. Chapman. Edited (for the Zoological Society
of London) by D. Sarr, M.A., F.R.S., F.Z.8., &c. London, 1896.
Price 10s.
The. Zoological Record, Volume the Thirty-third; being
Records of Zoological Literature relating chiefly to the year 1896.
By J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger,
W.A. Herdman, B. B. Woodward, A. W. Brown, D. Sharp, Florence
Buchanan, R. T. Giinther, and R. von Lendenfeld. Edited (for
the Zoological Society of London) by Davip Swarp, M.A., F.R.S.,
F.Z.8., &c. London, 1897. Price 30s.
The Zoological Record, Volume the Thirty-fourth; being
Records of Zoological Literature relating chiefly to the year 1897.
By J. A. Thomson, R. Lydekke1, R. Bowdler Sharpe, G. A. Boulenger,
W. A. Herdman, E. R. Sykes, E. A. Smith, G. C. Crick, A. W.
Brown, D. Sharp, F. A. Bather, Florence Buchanan, and R. yon
Lendenfeld. Edited (for the Zoological Society of London) by Davin
SHarp, M.A., F.R.S., F.Z.8., &. London, 1898. Price 30s.
The Zoological Record, Volume the Thirty-fifth ; being Records
ef Zoological Literature relating chiefly to the year 1898. By J. A.
Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger, W. A.
Herdman, EK. R. Sykes, E. A. Smith, G. C. Crick, A. W. Brown,
D. Sharp, F. A. Bather, Florence Buchanan, R. T. Giinther, and
R. yon Lendenfeld. Edited (for the Zoological Society of Londen) by
Davin Sarr, M.A., F.R.S., F.Z.8., &c. London, 1899. Price 30s.
The Zoological Record, Volume the Thirth-sixth ; being Records
of Zoological Literature relating chiefly to the year 1899. By
J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger,
W. A. Herdman, HK. R. Sykes, E. A. Smith, G. C. Crick, A. Willey,
A. W. Brown, D. Sharp, F. A. Bather, and R. von Lendenfeld.
Edited (for the Zoclogical Society of London) by Davip Suarp, M.A.,
F.RS., F.Z.8., &c. London, 1900. Price 30s.
: The Zoological Record, Volume the Thirty-seventh; being
Records of Zoological Literature relating chiefly to the year 1900.
By J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger,
W. A. Herdman, KE. R. Sykes, E. A. Smith, G. C. Crick, A. Willey,
A. W. Brown, D. Sharp, F. A. Bather, and E. A. Minchin. Edited
(for the Zoological Society of London) by Davin Sarr, M.A,,
E.R.S., F.Z.8., &e. London, 1901. Price 30s.
These publications may be obtained at the Socrmry’s Orrice
(3 Hanover Square, W.), of Messrs. Gurnry and Jackson (latei-
noster Row, H.C), or through any bookseller.
LIST OF INSTITUTIONS
TO WHICH
COPIES OF THE SOCIETY'S PUBLICATIONS ARE PRESENTED.
AFRICA,
The South-African Museum, Cape Town.
The South-African Philosophical Society, Cape Town.
The Museum, Durban, Natal.
AMERICA, SOUTH.
The National Museum, Buenos Ayres.
The Museum of Natural History, Santiago, Chili.
The Museum of La Plata, La Plata, Buenos Ayres.
AUSTRALASIA.
The Royal Society of Tasmania, Hobart.
The Royal Society of Victoria, Melbourne.
The Zoological and Acclimatization Society of Victoria, Melbourne,
The Linnean Society of New South Wales, Sydney.
The Royal Society of New South Wales, Sydney.
The New-Zealand Institute, Wellington.
AUSTRIA.
The Hungarian National Museum, Budapest.
The Imperial Academy of Sciences, Vienna.
The Zoological and Botanical Society, Vienna.
BELGIUM.
The Belgian Society of Geology, Paleontology and Hydrology,
Brussels.
The Congo Free State Museum, Teryueren, Brussels.
The Entomological Society of Belgium, Brussels.
The Malacological Society of Belgium, Brussels.
The Royal Academy of Sciences, Brussels.
The Royal Museum of Natural History, Brussels.
BRITISH INDIA.
The Asiatic Society of Bengal, Calcutta.
The Geological Survey of India, Calcutta.
The Indian Museum, Calcutta.
CANADA (DOMINION OF).
The McGill College, Montreal.
The Geological Survey of Canada, Ottawa.
The University of Toronto, Toronto,
2
CHINA.
The China Branch of the Royal Asiatic Society, Shanghai.
EAST INDIES.
The Royal Society of the Dutch East Indies, Batavia.
FRANCE.
The Linnean Society of Normandy, Caen.
The Agricultural Society, Lyons.
The Entomological Society of France, Paris.
The Museum of Natural History, Paris.
The National Society of Acclimatization, Paris.
The Zoological Society of France, Paris.
GERMANY.
The Royal Prussian Academy of Sciences, Berlin,
The Society of Friends of Natural History, Berlin.
The Natural-History Union for Rhineland and Westphalia, Bonn.
The Senckenbergian Society, Frankfort-on-Main.
The New Zoological Society, Frankfort-on-Main.
The Natural History Society, Freiburg-in-Breisgau.
The Royal Society of Sciences, Gottingen.
The Imperial Leopoldino-Carolinian Academy of Naturalists,
Halle.
The Natural-History Society, Halle.
The Natural-History Union, Hamburg.
The Royal Biological Station, Heligoland.
The Medical and Natural-History Society, Jena.
The Royal Bavarian Academy of Sciences, Munich.
The Union for Natural History of Wtrtemberg, Stuttgardt.
GREAT BRITAIN AND IRELAND.
The Belfast Natural History and Philosophical Society, Belfast.
The Philosophical Society, Cambridge.
The Royal Dublin Society, Dublin.
The Royal Irish Academy, Dublin.
The Royal Physical Society, Edinburgh.
The Royal Society, Edinburgh.
The Free Public Library and Museum, Liverpool.
The Athenzum Club, London.
The British Museum of Natural History, London.
The Entomological Society, London.
The Geological Society, London.
The King’s College Library, London.
The Linnean Society, London.
The London Institution.:
3
The Royal College of Physicians, London.
The Royal College of Surgeons, London.
The Royal Geographical Society, London.
The Royal Institution, London.
The Royal Society, London.
The University College, London.
The Literary and Philosophical Society, Manchester.
The Owens College, Manchester.
The Natural History Society, Newcastle-on-Tyne.
The Plymouth Institution and Devon and Cornwall Natural-History
Society, Plymouth.
The Marine Biological Laboratory, Plymouth.
The Yorkshire Philosophical Society, York.
HOLLAND.
The Royal Academy of Sciences, Amsterdam.
The Royal Zoological Society, Amsterdam.
The Dutch Society of Sciences, Haarlem.
The Dutch Entomological Union, The Hague.
The Royal Museum of the Netherlands, Leyden.
ITALY.
‘he Royal Institute of Superior Studies, Florence.
The Civil Museum of Natural History, Genoa.
The Italian Society of Natural Sciences, Milan.
The Zoological Station, Naples.
The Royal Academy of the Lincei, Rome.
The Royal Academy of Sciences, Turin.
JAPAN.
The Science College of the Imperial University, Tokyo.
RUSSIA.
The Society of Naturalists, Jurjeff (Dorpat).
The Society of Sciences of Finland, Helsingfors.
The Imperial Society of Naturalists, Moscow.
The Entomological Society of Russia, St. Petersburg.
The Imperial Academy of Sciences, St. Petersburg.
SCANDINAVIA.
The Bergen Museum, Bergen.
The Society of Sciences of Christiania, Christiania.
The Royal Danish Society of Sciences, Copenhagen.
The University Zoological Museum, Copenhagen.
The Royal Swedish Academy of Sciences, Stockholm.
The Royal Academy of Sciences, Upsala.
4
SPAIN, —
The Royal Academy of Sciences, Madrid.
SWITZERLAND.
The Philosophical and Natural-History Society, Geneva.
The Vaud Society of Natural Sciences, Lausanne.
The Society of Natural Sciences, Neuchatel.
The Natural-History Society, Zurich.
UNITED STATES OF AMERICA.
The Boston Society of Natural History, Boston.
The Museum of Comparative Zoology, Cambridge, Mass.
The Field Columbian Museum, Chicago.
The Illinois State Laboratory of Natural History, Illinois.
The American Journal of Science, Newhaven.
The American Museum of Natural History, New York.
The New-York Academy of Sciences, New York.
The Academy of Natural Sciences, Philadelphia.
The American Philosophical Society, Philadelphia.
The Entomological Society, Philadelphia.
The Essex Institute, Salem, Mass.
The Smithsonian Institution, Washington, D.C.
The United States Fish Commission, Washington, D.C.
The United-States Geological Survey, Washington, D.C.
The United-States National Museum, Washington, D.C.
WEST INDIES.
The Institute of Jamaica, Kingston.
LARADRALRARARAL PRPS
The Publications (except im special cases) are sent out direct as
soon as they are issued. It is requested that they may be ac-
knowledged by the return of the form of receipt sent with them,
in order that any mis-delivery may be brought to notice.
Publications sent in exchange to this Society should be addressed
to the Librarian at this Office. It is requested that they may be
sent direct by post, as much delay is caused by their transmission
through booksellers and in other ways.
By order of the Council,
P. L. SCLATER,
Secretary.
3 Hanover Sevare, Lonvon, W.,
August, 1902.
PROCEEDINGS
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
ZOOLOGICAL SOCIETY
OF LONDON,
1902, vol. I.
PART I.
CONTAINING PAPERS READ IN ~
JANUARY ann FEBRUARY..
JUNE ist, 1902.
PRINTED FOR THE SOCIETY,
SOLD AT THEIR HOUSE IN HANOVER SQUARE.
“LONDON:
MESSRS. L AGMANS, GREEN; AND CO.,
Ee =O
Prise tors" Sings a
LIST OF CONTENTS.
1902.—Vot. I.
Part | aaa ae,
January 14, 1902.
i Page
The Secretary. Report on the Additions to the Society's Menagerie in December 1901 .. Ay
Dr. A. 8. Woodward, F.R.8. Exhibition of a molar tooth of a Fossil Horse, Onohippidium. 1
Mr. Oldfield Thomas, F.R.S.. Exhibition of, and remarks upon, the skin and skull of a
Yellow-backed Duiker (Cephalophus syluiculiria) from N.H. Rhodesia
eee et oe eto ee
Mr. W. B. Tegetmeier, E.Z.8. Exhibition of the skin of a Mountain Hare eee
variabilis) which had been Bhated to belong to a Hare-Rabbit ce Minis cmt anh nen enen mee
1. On Waration in the Number and Arrangement of the ‘Male Genital Apertures in the
Norway Lobster (Nepinen te nor Wee By F. H. A. Marsuart, B.A., Christ’s Colles,
Cambridge ....-..++.....n0..-006 seeeteee sis ineieue Sinis's vivincield sie so e0is «eve wens sien 2
2. On some remarkable ee Adaptations in Dip uate has eee HINaR -
Lonnpera, C.M.Z.8 we epee ee eee ee cane san alsisicicleg ice sales ou sic m sole emanelnc aut ila
3. On the Specimen of the Quagga in the Imperial Museum of Natural History, Vienna.
By divpwie Vv. GuoREnz, CoML Zine. mates ss oleae ene nvamenistee etelcioiiols a ced Sea es 32
4. On a further Collection of Mammals made by Mr. Th. H. Lyle in Siam. By J. Lewis
Bonworn, MEAL. cesiciyosua cin Stepan etna ee oie ese erat acta eia terstee ah eee eee e eee “08
or
. On the Insects of the Order Rhynchota collected by Sir Harry Johnston, K.C.B., in the,
Uganda Protectorate. By W. L. DIsTanT .....+-+++20...seeees sireceere ate wennes 41
6. On two Collections of Lepidoptera made by Sir Harry Johnston, K.C.B., in the Uganda
Protectorate during the year 1900. By Arruur G. Borie, | Ph.D., F.LS., F.Z.8.;
&e. ; Senior SO eee ae laat Department, British Museum (Nat. Hist.)
(Plate I.).. eee CUP CRO BOC OI TRO ORO RO ICCCR BARRA ROCECe oy Gna . 44
February 4, 1902.
The Secretary. Report on the Additions to the Society's Menagerie in January 1902.... 51
Mr. F. E. Beddard, F.R.S. Exhibition of, and remarks upon, the malformed neck-
vertebrae of a Giraffe that had died in the Society’s Menagerie ........2+..++-+ s+ 52
Contents continued on page 3 of Wrapper.
ConTENTS (continued).
February 4, 1902 (continued).
Page
‘My. E. Degen. Notice of a Memoir on Ecdysis, as Morphological Evidence of the original
Tetradactyle Feathering of the Bird’s Fore-limb.......
Se i ce ee ee
ae, Notes on the Osteology of the Short-nosed Spoor W hale. By W. Buaxianp Bennam,
- D.Sc., M.A., F.Z.8., Professor of Biology in the es of punk New a aps
(Plates IT.-IV.).. a cama TS ele Safir aie Blwie le fc Synlaatuias’ ele aela eaters : coe
2, On a Collection of Dragonflies made re Members of the Skeat Expedition in the
- Malay Peninsula i in 1899-1900. By BF . Lamwuaw, B.A. (Plates.V.& VI.)......-. 68
3. List of a , small Collection of Orthopterous Insects formed by Sir Harry Johnston in
British Hast Africa and Uganda in 1899 and 1900, with Descriptions of five new
Species. By W. F. Kirsy, F.LS., F.H.S., Assistant in the Zoological Department,
British Museum (Natural History), ‘South Kensington afer st ialaitiu nt tase ay ae To ater an . 93
February 18, 1902.
Mr, L. W. aivie FE. Z.8. On the Identity of Lepidogaster stictopteryx Holt & Byrne with
DE MEE ROCEUNG TUB ETGOIE) oo sar etet tale aia telere Mca al wenen cro cle «Som tieisiavsacolalotele an eeiea. as 102
Mr. W. B. Tegetmeier, F.Z.8. Exhibition of, and remarks upon, the skull of a pecs
hybrid: between the Sheep and the Pig) 3/05). ee ee wees cece eles eneeerwe 102
Dr. C. I. Forsyth Major, F.Z.8. Exhibition of, and remarks upon, some jaws and teeth
of Pliocene Voles (Mimomys, gen. nov.) ......0+-+seeecaes Sinefetiis eOrecrutal sens stole rane 102
Mr. Lydekker. Exhibition of, and remarks upon, a skull and two pairs of antlers of an
Elk from Siberia
x
1. On Mustela paleattica from the Upper Miocene of Pikermi and Samos. By C. I.
Hageyrn ATOR, Helio (blate VIL.) ..).'. cclece tactic <eielatyeptanry ales Hobe DEE 109
2, On Two new Genera of Rodents from the Highlands of Bolivia. By Ouprretp Tuomas,
F.RS. (Plates VIII. & IX.) ..... nisjaiae sieyeienm thai ulein, watmin Seve apetalafcoureraiee hacen icine 114
3, On some New Mammals from Northern Nyasaland. By Ouprinytp Tuomas, F.R.S. .. 118
4. On some Characters distinguishing the Young of various Species of Polypterus. By
G, A. Bounzncmr, F.R.S. (Plates X. & XT.)......06..-.-.5.. isla erate iat eutaagehe ae eetee 121
_ 5. Description of a New Snake of the Genus Psammophis, from Cape Colony. By G. A.
Bovtmnemr, F.R.S. (Plate XIT.) ..............-. le spine AEROS he deena Ra He 126
6. Observations upon the Carpal Vibrisse in Mammals. By Frat E. pee M.A,
E.R. BA Vice-Secretary and Prosector of the Society ..-.-.eeee eee reccetenee cece 127°
1902— VOL. L
PART 1.04
Plate — oe 3
I. Lepidoptera from Uganda sve ngielenen ccimy ee meenntam
Il. : RO
IIT, ¢ Osteology of Cogia \... <6 vs ce cee sean ae cin A elt See ae
BY" Jae hia ean
V.
| Dragonflies from the Malay Peninsula «...--..2.20024.
VII. Skulls and Teeth of Mustela paleattica 2... ce. .eseee+ns
VIII. Neochowon Sioned oso sca ce witaiee + eg aow'e ane. eae ie
bea, + 052) abe Andinomys oda, adult ; ene i of Fa
4
X: 1,2. Polypterus lapradii. 3. P. weeksit ....6..s.cepsences
XI. 1. Polypterus congicus. 2. P. endlicheri. 3. P. senegalus
iad 4. DP. palmas ... eee ce ee ihe eee ee eee eee ne eee
XII. Psammophis leightoni Difonnaeu: fost dies aha ar
oy
The ‘ Proceedings’ for the year are issued in fowr parts, fommine, two volt rt
as follows :—
VOL, I.
Part I. containing papers read in J anuary and February, on June: st.
TH a e » March and April, on August Ist. a
VOL. II.
Part I. containing papers read in May and June, on Gotu Ist.
IL. 4s i », . November and December, on
PROCEEDINGS
_ GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
ZOOLOGICAL SOCIETY
OF LONDON
1902, vol. I.
PART) BE:
CONTAINING PAPERS READ IN
MARCH ann APRIL.
AUGUST ist, 1902.
PRINTED FOR THE SOCIETY,
SOLD AT THEIR HOUSE IN HANOVER SQUARE.
TON DON 5 sree |
MESSRS. LONGMANS DN AWD, 09.,
PATERN( aph-now 7 ob 0%& 4
SEP ZS | 902 fj
LIST OF CONTENTS.
1902.—Vot. I.
Part IT.
March 4, 1902.
Page
The Secretary. Report on the Additions to the Society’s Menagerie in February 1902.
(Plate RATT.) as esis ao's antares ateamebaioss thn parole meatsne eel ate plete ec taiec hae eon hee ra!
Mr. W. B. Tegetmeier, F.Z.S. Exhibition of a series of Poole ne of Prjevalsky’s Horse. 138
Mr. E. N. Buxton, F.Z.S, Exhibition of a series of Suey pe slides illustrative of Bird-
and Animal-life on the White Nile..-........... Roc OR pe mommies eee Nein Snes Willa
Mr. G. T. Bethune-Baker, ¥.Z.8. Notice of a Memoir on the hip potian oe of the f
Butterflies of the Family Lycenid@ ......00s. cece enc tees ee ances itera eee 88
1. On the Origin of Pearls. By H. Lystur Jamuson, M.A,, Ph.D. (Plates XIV.-XVII.).. 140 |
2, List of the Parrots represented in the Society’s Collection in January 1902, with Remarks
on some of the Rarer Species. By P. L. Sciarer, D.Sc., F.R.S., Secretary to the
Society.) (Bates XV TT ST ee creel areteiaietavere, eve icisse since’ es cfs atutcisl aipiaten aise eae 166
3. Descriptions of New Species of Coleoptera of the Family Halticide from South and
Central America. By Marvin Jacosy, F.H.S. (Plate XX.)........-...2-008 as ML
March 18, 1902.
Mr. Arthur Thomson. Report on the Insect-house for 1901 ......eeecscceeeeese sere 204
Mr. R. E. Holding. Exhibition of, and remarks upon, some malformed Horns and
Antlers: 2. 61 slot ele cicvsfasini icleisien'e plevevolerecaieie o letah wate aele| e/n atnl ain! ''s'9 wishallata\ecy (ries tata a aL
Lt.-Col. J. M. Fawcett. Notice of a Memoir on the Transformations of some South-African
Lepidoptera .......... SIONS TC HEE Se 6 BOO MO RE ia OICOGIAOIOn cH nolan cm OC iad! 205
1. The Evolution of Horns and Antlers. By Hans Ganpow, M.A., Ph.D., E.RS., F.Z.8... 206
2. On a new Stridulating-Organ in a Scorpion, By R. LE) Pocoon, FAS.) cc Ri teen. ome
%. On the Organ of Jacobson in the ae -Shrew Sas sae ela bs R.
Broom, M.D., B.Sc. (Plate XXI.).. SE aS ee salonid ap Urea:
Contents continued on page 3 of Wrapper,
~- ContTENTs (continued),
March 18, 1902 (continued).
Page.
4, On some Foraminifera and Ostracoda from Cocos Keeling Atoll, collected by Dr. C. W.
Andrews, 1898, By Frupericx Onapman, A.L.S., FARMS. «002-202 cece cere eee 228
5. Contributions to the Ichthyology of the Congo.—I. On some new Fishes from the -
French Congo. By G. A. Boutmnesr, F.R.S, (Plates XXIIXXIV.) ...,........ 234 -
April 15, 1902.
The Secretary, ue on the Additions to the eee 8 ae in ae 1902, *
(Plate XXV.).. ENE Sie OE EPI iS ie or Nismo ite yevies meee Oem
Prof. F. Jeffrey Bell, F.Z.S. Exhibition of, and remarks upon, a Starfish with injured
limbs which had undergone repair........- wang Sa Ntshe sosber cian ge shetty veers att 238
Dr, C. I. Forsyth Major, F.Z.S. Exhibition of, and remarks upon, some remains of a
co
| Piemy Hippopotamus from Cyprus ..-- +. se cece eee ee rece c cence cece cece tea g es 238
1, On the Windpipe and the Heart of the Condor. By Frank EH. Buppsrp. M.A., F.R.S., -
Vice-Secretary and) Prosector of the Society °.. 2. .ce ees egue ce cose Ouest nre vives te 239
2. On the Spiders of the Genus Latrodectus Walckenaer. By Freperick Pickarp Can-
BRIDGE, F.Z.S. (Plates XXVI. & XXVII.) ..........-6... Mie ne sipieaia cine pisickure din’ 247
. Notes on the Painted Snipe (Aostratula capensis) and Pheasant-tailed Jacana (Hydro-
phasianus chirurgus). By Frank Finn, B.A., F.Z.S., Deputy-Superintendent of the
tiidinmavisoum aOaleubta re comes cements x = alolc cise ceameanice tiles vieareroer nal dc 261
4. Contributions to the Ichthyology of the Congo,—II, On a Oollection of Fishes from
the Lindi River. By G. A. Bourencur, F.R.S, (Plates XXVIII.-XXX.)........., 265
5. Field-Notes upon some of the larger Mammals of Patagonia, made between September
1900 and June 1901. By Hueskeru PricnarD, F.Z.8. ae eee eect ee en ere mevee 272
6. Contributions to the Osteology of Birds.—Part V. Falconiformes. By. W. P. Pycrart,
Pais A, GS.ou(Plates XX XT ORM XTTT.) eae mele ein win ee poesoonnnen Pani ns 277
Index: cross ate 4 Sasac thon digi 6 Cet ese 55 os aH a EBOLOd Conan noc oowek avatsiven Lege
Titlepage «........-.. Pro eee peanaint eivie stern Seisteleaeictalal crapayelinlay «o/s!» ein’ dining oieleaiet eve ersveletalaneronees
PRD OM men Hal OURGCTE). ie iociee vie diate 4 see Sesie > miniaie aie visls eivin taicie o'sise/nioze veerees il
List of Contents ..... Sanaa at oie slolaparailvatns dia ve's MMB pis njnzue ee sve nies Stim eis eae ala gs ene vane lil
Alphabetical List of Contributors .......... 0s ss ccccesscsvaccens Panacea cael ix
List of Plates no Soc SAS UE TE Sela Genie Aer arc eanatsrsien petienateietie e xv
List of Text-figures .......:..+-..0- dee eee e eet eee nce et tee te eee RPE Ay istare SOWA
List of New Generic Terms .....0..00.0.eccseeecuees she eaehenel eorarerelte = Beene 5 une xix
=
LIST OF PLATES.
1902.—_V OL. I.
PART AE
Plate ay Page
DOLE Sf MUS PICUGIGNUE | iarniv'e6s Goals pea G Ade cate IPR AMT ES rea ea eter a 138
XIV.)
en I ameson on the Origin of Pearls ............s00eeeeececs 140
XV ES
XVILS i
XVIII. electus westermamnt, 3, QD crevececacneerersercececes :] 166
MK SPAN CercUs MAStETHAMUS 6. se nbc w aisle nse knead © ees |
XX. New Species of Halticide ....... raelah eee tide as Ete gre hae 171
XXI. Jacobson’s Organ in Macroscelides -.....-00sseeeeees a ove toe
XXII. 1. Allabenchelys longicuuda. 2. Clariallabes melas ....++..
EXD GeO! PURULE ao nk av gow whens ow 0’ craywhate eo BOeGH aie! caste pls eae mine eg |
DONTE SORMOCHTONIS CUPONEE wide an cosas veetarate st espe viriss Ses viola reapers
XXV. Cercopithecus otoleucus ©... scccoecccccscctccccscccenees Bag aes
es } Spiders of the Genus Latrodectus Sasatie a miastcn eye whee ea 247
XXVIII, 1, 2. Micralestes stormsi. 3. Phractura lindica ....... AD
XXIX. 1. Auchenoglanis punctatus. 2. Auchenoglanis pulcher, 3. Am-
philius DEVIS v.00 eee ee cece recent ence vente ee tees 265
XXX. I. Pseudoplesiops sguamiceps. 2. Tilapia stormsi. 3. Pris-
_ tigaster dollot ...... Mice ee Meech Severs Pana poe sic oe
XXXI. . if
XXXII. Osteology of the Falconiformes .....cecersseevccees ete NS Rhy ie
XXXIII.
NOTICE.
The * Proceedings’ for the year are issued in fowr parts, forming two volumes,
as followa:—
ye VOL. I.
Part I. containing papers read in January and February, on June Ist.
Ii. s ni » March and ae on ye Ist.
VOL. 1;
Part I. containing papers read in May and June, on October Ist.
If. 3, » November and December, on April Ist,
SER 9708
yar
Nae
Se
<
a5
yr
SMITHSONIAN INSTI
IANA ii
3 9088 00847 2185